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The Classification of Lower Organisms 

Ernst Hkinrich Haickei, in 1874 

From Rolschc (1906). 

By permission of Macrae Smith Company. 

C f 3 

The Classification 





PACIFIC ^., ^,kfi^..^ BOOKS 

Copyright 1956 by Herbert F. Copeland 
Library of Congress Catalog Card Number 56-7944 

Published by 


Palo Alto, California 

Printed and bound in the United States of America 


Chapter Page 

I. Introduction 1 

II. An Essay on Nomenclature 6 

III. Kingdom Mychota 12 

Phylum Archezoa 17 

Class 1. Schizophyta 18 

Order 1. Schizosporea 18 

Order 2. Actinomycetalea 24 

Order 3. Caulobacterialea 25 

Class 2. Myxoschizomycetes 27 

Order 1. Myxobactralea 27 

Order 2. Spirochaetalea 28 

Class 3. Archiplastidea 29 

Order 1. Rhodobacteria 31 

Order 2. Sphaerotilalea 33 

Order 3. Coccogonea 33 

Order 4. Gloiophycea 33 

IV. Kingdom Protoctista 37 

V. Phylum Rhodophyta 40 

Class 1. Bangialea 41 

Order Bangiacea 41 

Class 2. Heterocarpea 44 

Order 1. Cryptospermea 47 

Order 2. Sphaerococcoidea 47 

Order 3. Gelidialea 49 

Order 4. Furccllariea 50 

Order 5. Coeloblastea 51 

Order 6. Floridea 51 

VI. Phylum Phaeophyta 53 

Class 1. Heterokonta 55 

Order 1. Ochromonadalea 57 

Order 2. Silicoflagellata 61 

Order 3. Vaucheriacea 63 

Order 4. Choanoflagellata 67 

Order 5. Hyphochytrialea 69 

Class 2. Bacillariacea 69 

Order 1. Disciformia 73 

Order 2. Diatomea 74 

Class 3. Oomycetes 76 

Order 1. Saprolegnina 77 

Order 2. Peronosporina 80 

Order 3. Lagenidialea 81 

Class 4. Melanophycea 82 

Order 1 . Phaeozoosporea 86 

Order 2. Sphacelarialea 86 

Order 3. Dictyotea 86 

Order 4. Sporochnoidea 87 

V ly 

Chapter Page 

Orders. Cutlerialea 88 

Order 6. Laminariea 89 

Order 7. Fucoidea 91 

VII. Phylum Pyrrhophyta 94 

Class Mastigophora 95 

Order 1. Cryptomonadalea 96 

Order 2. Adiniferidea 98 

Order 3. Cystoflagellata 99 

Order 4. Cilioflagellata 102 

Order 5. Astoma 105 

VIII. Phylum Opisthokonta 110 

Class Archimycetes Ill 

Order 1. Monoblepharidalea Ill 

Order 2. Chytridinea 113 

IX. Phylum Inophyta 119 

Class 1. Zygomycetes 121 

Order 1. Mucorina 121 

Order 2. Entomophthorinea 124 

Class 2. Ascomycetes 125 

Order 1. Endomycetalea 129 

Order 2. Mucedines 130 

Order 3. Perisporiacea 131 

Order 4. Phacidialea 133 

Order 5. Cupulata 134 

Order 6. Exoascalea 137 

Order 7. Sclerocarpa 137 

Order 8. Laboulbenialea 140 

Class 3. Hyphomycetes 140 

Order 1. Phomatalea .... 141 

Order 2. Melanconialea 141 

Order 3. Nematothecia 141 

Class 4. Basidiomycetes 142 

Order 1. Protobasidiomycetes 146 

Order 2. Hypodermia 147 

Order 3. Ustilaginea 149 

Order 4. Tremcllina 149 

Order 5. Dacryomycetalea 150 

Order 6. Fungi 150 

Order 7. Dermatocarpa 152 

X. Phylum Protoplasta 157 

Class 1. Zoomastigoda 157 

Order 1. Rhizoflagellata 158 

Order 2. Polymastigida 163 

Order 3. Trichomonadina 166 

Order 4. Hypcrmastiglna 168 

Class 2. Mycetozoa 171 

Order 1. Enteridiea 171 

Order 2. Exosporea 177 


Chapter Page 

Order 3. Phytomyxida 177 

Class 3. Rhizopoda 179 

Order 1. Monosomatia 183 

Order 2. Miliolidea 185 

Order 3. Foraminifera . . . 185 

Order 4. Globigerinidea 187 

Order 5. Nummulidnidea 188 

Class 4. Heliozoa 189 

Order 1. Radioflagellata 190 

Order 2. Radiolaria 194 

Order 3. Acantharia 195 

Order 4. Monopylaria 198 

Orders. Phaeosphaeria 198 

Class 5. Sarkodina 200 

Order 1. Nuda 201 

Order 2. Lampramoebae 205 

XI. Phylum Fungilli 206 

Class 1. Sporozoa 207 

Order 1. Oligosporea 209 

Order 2. Polysporea 211 

Order 3. Gymnosporidiida 211 

Order 4. Dolichocystida 214 

Orders. Schizogregarinida 215 

Order 6. Monocystidea 215 

Order 7. Polycystidea 216 

Order 8. Haplosporidiidea 218 

Class 2. Neosporidia 219 

Order 1. Phaenocystes 219 

Order 2. Actinomyxida 221 

Order 3. Cryptocystes 222 

XII. Phylum Ciliophora 223 

Class 1. Infusoria 228 

Order 1. Opalinalea 228 

Order 2. Holotricha 229 

Order 3. Heterotricha 230 

Order 4. Hypotricha 233 

Order 5. Stomatoda 233 

Class 2. Tentaculifera 235 

Order Suctoria 235 

List of Nomenclatural Novelties 237 

Bibliography 238 

Index 271 



Portrait of Ernst Heinrich Haeckel Frontispiece 

Figure Page 

1. Structure of cells of blue-green algae 13 

2. Photographs of Escherichia coli . . . 15 

3. Caulobacterialea; Myxobactralea; Cristispira Veneris 26 

4. Coccogonea; Gloiophycea 32 

5. Bangialea 42 

6. Nuclear phenomena in Polysiphonia violacea 45 

7. Heterocarpea 48 

8. Ochromonadalea 54 

9. Ochromonadalea; Silicoflagellata 56 

10. Vaucheriacea 64 

11. Choanoflagellata 68 

12. Hyphochytrialea 70 

13. Bacillariacea 72 

14. Oomycetes 78 

15. Stages of nuclear division in Stypocaulon 84 

16. Familiar kelps of Pacific North America 90 

17. Microscopic reproductive structures of Laminaria yezoensis ... 92 

18. Cryptomonadalea 97 

19. Cystoflagellata; Cilioflagellata 104 

20. Astoma 106 

21. Astoma 108 

22. Monoblepharidalea 114 

23. Chytridinea 116 

24. Zygomycetes 122 

25. Ascomycetes 132 

26. Ascomycetes 136 

27. Mycosphaerella personata 138 

28. Basidiomycetes 144 

29. Fruits of Agaricacea 153 

30. Rhizoflagellata 160 

31. Polymastigida; Trichomonadina 164 

32. Hypermastigina 170 

33. Mycetozoa 176 

34. Ceratiomyxafruticulosa 178 

35. Life cycle of "Tretomphalus" i. e., Discorbis or Cymbalo por a . . . 180 

36. Shells of Rhizopoda 184 

37. Radioflagellata 192 

38. Radiolaria; Acantharia; Monopylaria; Phaeosphaeria 196 

39. Chaos Protheus 200 

40. Sarkodina 204 

41. Life cycle of Goussia Schuhergi 208 

42. LUe cycle of Plasmodium; Babesia bigemina 212 

43. Life cycle of Myxoceros Blennius 220 

44. Infusoria, order Hypotricha 232 

45. Tokophrya Lemnarum 234 


Chapter I 

The purpose of this work is to persuade the community of biologists that the ac- 
cepted primary classification of living things as two kingdoms, plants and animals, 
should be abandoned; that the kingdoms of plants and animals are to be given definite 
limits, and that the organisms excluded from them are to be organized as two other 
kingdoms. The names of the additional kingdoms, as fixed by generally accepted 
principles of nomenclature, appear to be respectively Mychota and Protoctista. 

These ideas originated, so far as I am concerned, in the instruction of Edwin 
Bingham Copeland, my father, who, when I was scarcely of high school age, admitted 
me to his college course in elementary botany. He thought it right to teach freshmen 
the fundamental principles of classification. These include the following: 

The kinds of organisms constitute a system of groups; the groups and the system 
exist in nature, and are to be discovered by man, not devised or constructed. The 
system is of a definite and peculiar pattern. By every feature of this pattern, we are 
inductively convinced that the kinds of organisms, the groups, and the system are 
products of evolution. It is this system that is properly designated the natural system 
or the natural classification of organisms. It is only by metaphor or ellipsis that these 
terms can be applied to systems formulated by men and published in books. 

Men have developed a classification of organisms which may be called the taxo- 
nomic system. Its function — the purpose for which men have constructed it — is to 
serve as an index to all that is known about organisms. This system is subject to cer- 
tain conventions which experience has shown to be expedient. Among natural groups, 
there are intergradations; taxonomic groups are conceived as sharply limited. Natural 
groups are not of definite grades; taxonomic groups are assigned to grades. When we 
say that Pisces and Filicineae are classes, we are expressing a fact of human conven- 
ience, not a fact of nature. The names assigned to groups are obviously conventional. 

Since the taxonomic system represents knowledge, and since knowledge is ad- 
vancing, this system is inherently subject to change. It is the right and duty of every 
person who thinks that the taxonomic system can be improved to propose to change 
it. A salutary convention requires that proposals in taxonomy be unequivocal: one 
proposes a change by publishing it as in effect; it comes actually into effect in the 
degree that the generality of students of classification accept it. The changes which 
are accepted are those which appear to make the taxonomic system, within its conven- 
tions, a better representation of the natural system. Different presentations of the 
taxonomic system are related to the natural system as pictures of a tree, by artists of 
different degrees of skill or of different schools, are related to the actual tree; the 
taxonomic system is a conventionalized representation of the natural system so far as 
the natural system is known. 

These statements are intended to make several points. First, as a personal matter, 
advancement of knowledge of natural classification, and corresponding improvement 
of the taxonomic system, have been my purpose during the greater part of a normal 
lifetime. Secondly, I have pursued this purpose, and continue to pursue it, under the 
guidance of principles which all students of classification will accept (perhaps with 
variations in the words in which they are stated). In the third place, I have tried to 
answer the question which scientists other than students of classification, and likewise 
the laity, are always asking us: why can one not leave accepted classification undis- 

2 ] The Classification of Lower Organisms 

turbed? One proposes changes in order to express what one supposes to be improved 
knowledge of the kinds of organisms which belong together as facts of nature. If here 
I place bacteria in a different kingdom from plants, and Infusoria in a different king- 
dom from animals, it is because I believe that everyone will have a better understand- 
ing of each of these four groups if he does not think of any two of them as belonging 
to the same kingdom. 

The course of evolution believed to have produced those features of the natural 
system to which the present work gives taxonomic expression is next to be described. 

Life originated on this earth, by natural processes, under conditions other than 
those of the present, once only. These are the opinions of Oparin ( 1938) 1, and appear 
sound, although some of the details which he suggested may not be. When the crust 
of the earth first became cool, it was covered by an atmosphere of ammonia, water 
vapor, and methane, and by an ocean containing the gases in the atmosphere above 
it and minerals dissolved from the crust. This is to state the hypotheses that organic 
matter in the form of methane is older than life; and that whereas conditions on the 
face of the earth tend now to cause oxidation, they tended originally to cause reduc- 
tion. In a medium of the nature of the supposed primitive ocean, spontanous chemical 
changes will occur and produce organic compounds of considerable complexity: this 
has repeatedly been demonstrated by experiment. To convert a solution of ammonia, 
methane, and minerals into protoplasm, Oparin postulates a very long series of 
changes, producing successively more complicated compounds and mixtures, and re- 
quiring perhaps hundreds of millions of years. The changes are conceived as acci- 
dents; they are supposed to have been probable accidents, like throwing a seven at dice, 
not events which could only very rarely occur by accident, like throwing twenty sevens 
in succession. By supposing that some of these processes used up the m.aterials neces- 
sary for them, Oparin provides an explanation of the single origin of life: we are 
confident that all life is of one origin, because all protoplasm is of the same general 
nature, and all life consists of essentially the same processes. The course of events 
described would have yielded, as the original form of life, anaerobic saprophytes; this 
is in harmony with the fact that anaerobic energesis is in a sense the basic metabolic 
process. The original organisms would scarcely have possessed nuclei: Oparin's 
theories indicate, as the most primitive form of life which has been able to survive, 
the anaerobic bacteria. The anaerobic bacteria are indeed very far removed from any 
lifeless things; their protoplasm and their metabolism are fundamentally the same 
as ours. 

Life requires energy. Under anaerobic conditions, an organism can obtain energy 
by converting sugars to alcohol, but it can not use alcohol as a source of energy. This 
example means that anaerobic energesis yields energy in strictly limited quantity and 
produces incompletely oxidized compounds. So long as all life was anaerobic, it was 
engaged in converting the organic matter upon which it depended into forms which 
it could not use; life under these conditions, at least if they persisted for any great 
period of time, was surely very sluggish. A further scries of changes in the metabolic 
system, occurring accidentally in certain organisms and preserved by natural selec- 
tion, brought photosynthesis into existence. The purple bacteria are believed to rep- 
resent stages in the evolution of photosynthesis, which exists in its fully developed 
form, involving the release of elemental oxygen, in the blue-green algae. Once photo- 

^ Dates in parentheses are references to works which have been consulted and listed in 
the bibliography. 

Introduction [ 3 

synthesis was established in certain organisms, aerobic energesis became possible both 
to these and to others. This made possible a manner of life more vigorously active 
than before. The inconsiderable groups of autotrophic bacteria — the organisms which 
live by oxidizing inorganic matter — appear to be secondary developments dependent 
upon the existence of photosynthesis. 

The organisms whose origin has been suggested thus far — the ordinary bacteria, 
anaerobic and aerobic, the autotrophic bacteria, the purple bacteria, and the blue- 
green algae — are relatively simple in structure and function; all consist of minute 
physiologically independent cells. The first step in the evolution of more complex 
organisms was the evolution of the nucleus. 

Morphologically, the nucleus is a part of a protoplast which is set apart by a mem- 
brane and which originates ordinarily by division of a pre-existent nucleus in the 
manner called mitosis. In this process, a definite number of definite chromosomes 
appear and undergo equal division. The nucleus exercises control over the protoplast 
in which it lies. Its controlling action depends upon the chromosomes which go into 
it, and mitosis has the effect that all nuclei which are derived from one original nu- 
cleus strictly by normal processes of mitosis are identical in the controlling effects 
which they exert. Thus the nucleus serves for the precise transmission of a compli- 
cated heredity. Beside mitosis, there are two other processes — two only — meiosis and 
karyogamy, by which nuclei may produce other normal and enduringly viable nuclei. 
In a sequence of generations of individuals sexually produced, these processes occur 
alternately, each one at one point in each cycle of sexual i-eproductlon. Mendelian 
heredity is produced by changes, in the sets of chromosomes (or parts of chromo- 
somes) in individual nuclei, which occur during meiosis and karyogamy. The role of 
the nucleus in sexual reproduction is one of its essential characters: the nucleus is re- 
lated to sexual reproduction, including Mendelian heredity, as structure to function. 

The existence of organisms without nuclei shows that the nucleus evolved after life 
did: it did not evolve at the same time as protoplasm. The essential uniformity of 
the nucleus and of its association with sexual reproduction shows that these things 
evolved only once, and together. There are a very few organisms, as Porphyridium 
and Prasiola, in which the presence or absence of nuclei is not certain; there is ac- 
cordingly scant evidence for speculation as to the manner of this evolution. As to the 
tim.e, we know only that microfossils representing nucleate organisms occur in the 
uppermost strata of the Proterozoic era. 

By making possible the precise transmission of a complicated heredity, the nucleus 
has made possible the development of complexities of structure and function exceed- 
ing by far anything occurring in non-nucleate organisms. It appears that as soon as 
the nucleus was in existence, organisms provided with it entered upon evolution in 
many characters and gave rise to many distinguishable groups. Among these groups, 
those which consist respectively of the typical plants and the typical animals are the 
greatest. There is, however, neither any a priori reason, nor any evidence from nature, 
for a belief that all groups of nucleate organisms must naturally belong to one or the 
other of these two. Several other groups, in general much less considerable than these, 
are thoroughly distinct and appear equally ancient. 

E. B. Copeland understood the history of life very much as it has just been pre- 
sented. In his teaching, he treated the bacteria and blue-green algae as standing en- 
tirely apart both from plants and from animals, and pointed out several other groups 
which are not as a matter of nature either plants or animals. It was his opinion that 
these groups should be treated as a series of minor kingdoms; he excused himself 

4 ] The Classification of Lower Organisms 

from the attempt to formulate a definite and comprehensive system. This teaching 
was the original stimulus which has led to the present work. I bear witness that E. B. 
Copeland taught these things in 1914; he did not publish them until he had ceased 
to teach (1927). 

In the year 1926, when the teaching of elementary botany was first fully my own 
responsibility, I came to the conclusion that the establishment of several kingdoms 
of nucleate organisms in addition to plants and animals is not feasible; that all of 
these organisms are to be treated as one kingdom. This is one of the few points of 
originality which I claim for my work. It is true that the kingdom thus described is 
not very different from the third kingdom which various early authors proposed and 
which Haeckel (1866) named Protista. Haeckel, however, in his varied presentations 
of the kingdom Protista, included always the bacteria. By setting apart the bacteria 
and blue-green algae as yet another kingdom, one meets, at least in part, the objection 
to the "third kingdom" that it is heterogeneous beyond what can be tolerated. 

It has been necessary to meet also the objection that the "third kingdom" substi- 
tutes, for an acknowledgedly vague boundary between plants and animals, two vague 
boundaries: it has been necessary to recognize characters by which sharp definition 
can be given to plants and animals. It is my contention that these characters have 
long been known. The kingdom of plants, as the taxonomic representation of a 
natural group, is to be defined by the system of chloroplast pigments described by 
Willstatter and Stoll (1913), and also by the production of certain carbohydrates 
which occur only sporadically elsewhere. The kingdom of animals is defined by em- 
bryonic development through the stages called blastula and gastrula, as pointed out 
by Haeckel (1872). It is believed that no organisms exhibit both of these sets of 
characters; the "third kingdom" includes the nucleate organisms which exhibit 
neither. The kingdoms of plants and animals as here defined are essentially those 
which are traditionally and popularly accepted. They include all the creatures which 
Linnaeus listed as plants and animals, with the exceptions of forms of which he knew 
little, and which he listed superficially at the ends of his treatments of the respective 

Of course, the definitions are not warranted to describe the kingdoms without ex- 
ception. For one thing, each is supposed to have come into existence by evolution 
through a line of organisms which exhibited its characters imperfectly. For another, 
evolution can erase what it has created; it is proper to include in a group organisms 
which have by degeneration lost its formal characters. These things are true of all 
taxonomic groups. 

In due form, then, the system of kingdoms here maintained is as follows: 

Kingdom I. Mychota. Organisms without nuclei; the bacteria and blue-green 

Kingdom II. Protoctista. Nucleate organisms not of the characters of plants and 
animals; the protozoa, the red and brown algae, and the fungi. 

Kingdom III. Plantae. Organisms in whose cells occur chloroplasts, being plastids 
of a bright green color, containing the pigments chlorophyll a, chlorophyll h, carotin, 
and xanthophyll, and no others; and which produce sucrose, true starch, and true 

Kingdom IV. Animalia. Multicellular organisms which pass during development 
through the stages called blastula and gastrula; typically predatory, and accordingly 
consisting of unwalled cells and attaining high complexity of structure and function. 

This system has twice been given brief publication (1938, 1947). I am glad to say 

Introduction [ 5 

that Barkley (1939, 1949) and Rothmaler (1948) maintain a system of kingdoms 
which differs from this in a single significant detail. 

Assuming that this system is tenable as a matter of reason, it will nevertheless not 
be accepted among taxonomists unless they have some knowledge of what it means 
in detail. No person is called upon to recognize the kingdoms Mychota and Protoc- 
tista until systems of their subordinate groups are available. The bulk of the present 
work consists of such systems. Complete systems of divisions or phyla, classes, and 
orders are presented. Groups of lower rank are presented in part, as examples. As a 
matter of facility, the groups of lower rank are presented more fully in the smaller or 
better known groups than in the larger or more obscure. 

The preparation of this work has taken more than ten years. In the course of it I 
have received much help. Among those who have answered queries, or who have in 
various drafts scrutinized the whole work or parts of it for faults of every degree of 
significance, are Dr. G. M. Smith of Stanford University; Dr. A. S. Campbell of St. 
Mary's College; Dr. Herbert Graham, formerly of Mills College; Dr. Lee Bonar, Dr. 
G. L. Papenfuss, and Dr. H. L. Mason of the University of California at Berkeley; 
Dr. E. R. Noble of the University of California at Santa Barbara; and Dr. H. C. Day 
of Sacramento Junior College. The counsel of E. B. Copeland has not been withheld. 
It is a matter of grief that two distinguished zoologists of the University of California, 
Dr. S. F. Light and Dr. Harold Kirby, have passed away during the long course of 
this work; as have two colleagues who were my closest friends, Dr. H. J. Child and 
Dr. C. C. Wright. 

The portrait of Haeckel which is my frontispiece is used by permission of Macrae 
Smith Company, Philadelphia. Two figures of Chrysocapsa are used by permission 
of the Cambridge University Press. Numerous figures have been taken from the 
Archiv filr Protistenkunde with the gracious permission of Prof. Dr. Max Hartmann. 

We do well to realize our indebtedness to libraries and librarians. To a great extent, 
this work has been made possible by the unstinted hospitality of the Biology Library 
of the University of California at Berkeley. 

Two statements appear regularly in prefaces; they are of truths which are strongly 
impressed upon authors. In the first place, those who have given help have made the 
work better; the author alone is responsible for deficiencies. The foregoing list of 
good friends and good scholars does not claim them as proponents of the thesis of 
this work. 

In the second place, the work is not offered as perfect or nearly so. The scholar in 
a strictly limited field may become master of the available knowledge. One who at- 
tempts studies in a broad field realizes that he is dealing with many subjects of which 
others know far more than he; that he has not wrung dry the existing literature; that 
some of the problems which puzzle him will be solved if he will wait a little longer. 
His colleagues have a right to raise these matters as criticisms. But surely, it is not 
desired that studies in broad fields be never attempted or indefinitely delayed. 

A matter which is particularly likely to arouse criticism is that of the names which 
are here applied to the groups. The principles according to which this has been done 
are set forth in the following chapter. I beg my colleagues, in dealing with this chapter 
and with the names subsequently applied, not to imagine that I have acted without 
grave thought. I have decided, that as in classification, so also in nomenclature, I 
should set before the community of biologists an experiment in the application of 
principles; among which principles there are surely some whose strict application 
will be to the good of our science. 

Chapter II 


Whoever sets forth a system of groups finds himself under the necessity of making 
responsible decisions as to names. The kingdoms have received more names than one 
(Table 1 ), and so have nearly all of the major groups within them: it has here been 
necessary to decide as to the validity and application of the names Flagellata and 
Mastigophora, Rhodophyceae and Florideae, Rhizopoda and Sarcodina, and many 

TABLE 1. Names Applied by Various Authors to the Kingdoms 
OF Systems of Four Kingdoms 




1938, and 



Haeckel, 1894 

Barkley, 1939 

1948 1947 and here 

I Protophyta 




II Protozoa 




III Metaphyta 




IV Metazoa 




In dealing with plants, with animals, or with bacteria, it is necessary to observe 
the codes of nomenclature enacted by international congresses for the respective 
groups: the botanical code (Fournier, 1867; Lanjouw, 1952), with amendments 
enacted in 1954; the zoological code of 1889 as amended in 1948 and 1953 (issue of 
an edition incorporating the amendments is expected; Hemming, 1954); and the 
bacteriological code (Buchanan et al., 1948). Breach of the appropriate code renders 
an author liable to the penalty of having his work treated as nullity. 

The existence of three sets of rules for one thing, and the continual amendment of 
the older codes, are evidence of imperfection. It will not be purely destructive to 
point out certain anomalies in the codes as they stand. 

The zoological code pretends to overrule the principles of grammar in treating 
specific epithets as names. It is true that some of these words are names: the Catus in 
Felis Catus is a name of the cat, and the Mays in Zea Mays is a name of maize. But 
the great majority are adjectives; the sapiens in Homo sapiens is not by itself a des- 
ignation of man, and the vulgarc in Hordeum vulgarc is not a name of barley. It is a 
further offense against grammar that the code prescribes, as the names of all families 
of animals, adjectives in the feminine. Applied originally to families of birds, Aves, 
these names were unobjectionable; but the names of the kingdom and of the over- 
whelming majority of its subordinate groups are neuter. 

The botanical code as published with its appendages makes a book of more than 
two hundred pages. A statement of principles, in which the last clause provides for 
exceptions, occupies two pages. The definite rules and recommendations occupy 
about thirty-five pages; one who studies them critically will find that they prescribe 
more than one procedure not warranted by principle. A list of names maintained or 
rejected irrespective of principle occupies about seventy pages. These things mean 
that current botanical nomenclature is only within limits a matter of rule; it is to a 
considerable extent governed by enactments of the nature of ex post facto laws and 
bills of attainder. 

An Essay on Nomenclature [ 7 

The bacteriological code is for the most part a condensation of an earlier edition 
of the botanical code. It includes the odd feature that the name of a genus of bacteria 
is to be changed if it had previously been used either among plants or among Protozoa. 
Since there is an earlier Phytomonas among flagellates, bacteriologists have given a 
new name to the bacterium Phytomonas. The avoidance of homonyms which they 
desire will not, however, be attained: no zoologist will allow a new name for the 
flagellate Klebsiella on account of an earlier Klebsiella among bacteria. 

The grounds upon which these things are treated as wrong are provided by a 
passage in the botanical laws of 1867 which is believed to define the legitimate 
authority of congresses and codes: 

"Les regies de la nomenclature ne pouvent etre ni arbitraires ni imposees. Elles 
doivent etre bassees sur des motifs assez clairs et assez forts pour que chacun les 
comprenne et soit dispose a les accepter." 

It is implied by this statement that principles, appealing to the reason and found 
sound by the trial of experience, were in existence when it was written; and this is 
the truth. By this statement, the legitimate powers of congresses are those of courts 
of common law, which avoid the explicit making of law, but discover the law, inter- 
pret it, and apply it. Congresses and codes may legitimately (a) state explicitly 
corollaries of the principles when they are not obvious; and (b) determine arbitrarily 
matters which are necessarily determined arbitrarily, not being within the range of 
principle. One would not in theory deny a power (c) to validate breaches of principle 
when these are of an expedience verging on necessity; but its use by botanical con- 
gresses to produce a roll of exceptions of twice the bulk of the text of the code leads 
one to doubt the expedience of this admission. It has been through failure to recog- 
nize the legitimate limits of their powers — through a conception that their powers 
are sovereign or plenary — that international congresses have come to enact codes 
conflicting with each other and giving incomplete satisfaction in themselves. 

Under these circumstances, a nomenclature of superior legitimacy can be applied 
in groups treated as removed from the jurisdiction of the codes. Not without diffi- 
dence, this assumption is extended to the bacteria; it will be agreed that the nomen- 
clatural practice applied to the bacteria must be the same as that which is applied 
to the blue-green algae. 

Here one attempts a brief formulation of those principles, appealing to reason 
and proven sound in practice, to which all nomenclature must conform. 

1. Scientific names are words of the Latin language. They are not "of Latin form" 
or "construed as Latin"; they are Latin. This is to treat Latin as a living language and 
scientific names as subject to the rules of its grammar. They are not code-designa- 
tions, nor words of any language or none, as chemical names are. 

2. The name of a group of the kind called a genus is a proper noun in the singular. 
Linnaeus replaced all generic names which were adjectives; all of us his successors 
should do likewise. 

3. The names of groups of genera are proper nouns, or adjectives used as proper 
nouns, in the plural. 

The foregoing principles are of pre-Linnaean origin; beginning with his first sig- 
nificant work (1735), Linnaeus took them for granted. For the principle next to be 
stated, authority is the practice of Linnaeus in later works (1753 and subsequently) : 

4. The name of a species consists of the name of the genus to which it belongs fol- 
lowed by one epithet, ordinarily an adjective, occasionally a noun in apposition or 
in the genitive. 

8 ] The Classification of Lower Organisms 

A fifth principle represents Linnaean practices as subsequently modified: 

5. Named taxonomic groups are necessarily of certain fixed ranks called categories, 
i.e., lists. There are seven principal categories, specified as follows. Every individual 
organism belongs to a group conceived as the single kind and called a species. Every 
species belongs to a genus; every genus to a family; every family to an order; every 
order to a class; every class to a division or phylum; ever)' division or phylum to a 
kingdom. These conventions have the effect that the groups of each principal category 
embrace the entire range of the kinds of organisms. 

The categories of genera and species come down from classic antiquity. Linnaeus 
originated orders; he originated classes in the sense of named definite groups; and it 
appears that he is responsible for kingdoms: the writer knows of no earlier authority 
for the traditional three kingdoms of nature. The category next below that of king- 
doms has been variously called; originally it was emhranchements (Cuvier, 1812). 
The history of the category of families is somewhat involved. It originated in the 
work of Adanson (1763); in the following year, Linnaeus (1764) treated the groups 
which Adanson had called families as natural orders. Botanists for a long time held 
that families and orders are the same thing. Zoological practice gradually made fam- 
ilies a separate category. Authority for the list of seven principal categories as given 
is Agassiz (1857). 

Nothing prevents the assignment of groups to categories other than these, to sub- 
classes, tribes, and the like. These may be called subordinate categories. The groups 
of any subordinate category embrace only fragments of the range of kinds of 

The work of Linnaeus was largely innovation, and he did not have the face to de- 
clare binding the generally accepted rule of priority. Definite authority for the rule 
is de Candolle (1813). As currently applied, it may be stated as follows: 

6. The valid name of a group is its oldest published name, conforming to the rules, 
and not previously applied in the same kingdom. 

As corollaries of the rule of priority, when groups are combined, the oldest name 
of any of them must be applied to the whole, and when a group is divided, its name 
must be retained for one of the parts. The part to which the original name is to be 
applied is determined by the method of types, formulated by Strickland and his as- 
sociates (1843) : 

7. When a group is divided, its name must be applied to the portion which includes 
whatever part of it the original author would have regarded as typical. The part thus 
specified is the nomcnclatural type of the group. 

In the application of these principles to the naming of the groups of Mychota and 
Protoctista, the following practices appear expedient. 

A name is applied by publication in such fashion that the community of biologists 
may reasonably be held responsible for knowing of its existence and recognizing the 
entity to which it is to be applied. This means that it is to be printed in a technical 
book or journal and defined in a language for which the generality of biologists will 
not require an interpreter, namely Latin, English, French, or German. Any regulation 
more detailed than this is an excuse for breaches of priority. Definition is not neces- 
sarily by description: nearly all of the Linnaean genera of plants were established 
by the listing of species in the Species Plantarum. 

When two or more groups published in the same work at the same time are to be 
combined, their names are of equal priority. The choice of one of their names by the 
first author who combines them is binding. 

An Essay on Nomenclature [ 9 

A type as specified in the original publication of a group, or as implied by the in- 
clusion of a single subordinate group, is unchangeable. Linnaeus and his immediate 
successors had no conception of the device of types, and it is practically impossible 
to be certain of the elements which they would have regarded as typical in some of 
their groups. It remains necessary that the type system be applied to these groups. In 
some of them, it may be expedient that international authority, proceeding with due 
caution, declare types arbitrarily. An individual scholar will do better to call what he 
supposes to be the type of a group by a difTerent term, namely standard (Sprague, 
1926) : the standard of a group is a supposed type which remains open to debate. The 
framers of codes have undertaken to make binding the choice of a type by the first 
author who divides a group. On various occasions, however, this action has been 
demonstrably mistaken. 

Certain venerable names, as Vermes and Algae as used by Linnaeus, were applied 
to altogether miscellaneous collections of organisms among which the selection of a 
standard would be purely arbitrary. Such names are called nomina confusa, and are 
to be abandoned. 

It follows from the principle of the binomial nomenclature of species that no genus 
is named until one or more of its species are designated by binomial names. It fol- 
lows also that in works in which the nomenclature of species is not definitely binomial 
no names are of any standing. Hence, the point of time from which priority is effective 
is that of the introduction of binomial nomenclature, namely 1753. The enactment of 
other starting points for the nomenclature of particular groups is pretended law 
which is not law, like the pretended laws of American states which attempt to regu- 
late interstate commerce under the appearance of doing something else. 

The original spelling of names, so far as it is tolerable Latin, is not to be changed. 
Errors of gender or number, obvious mistakes of spelling, and misprints, are to be 
corrected. Good Latin is written without diacritical marks: a German Umlaut in a 
name as published is corrected by inserting an e; accents, cedilles, and other barbar- 
isms are dropped. The codes err in prescribing changes in spelling beyond those 
which are here admitted. If they should establish uniformity in the future, it would 
be at the expense of divergence from the most respected works of the past. 

Specific epithets are capitalized if they are ( 1 ) names in the nominative, in ap- 
position with the generic names; (2) names of persons, places, or organisms in the 
genitive; (3) adjectives derived from names of persons. 

Transfer of groups from one kingdom to another does not warrant any meddling 
with names. When a group is transferred from one kingdom to another, its valid name 
in the former — its oldest name not previously used in the kingdom in which it was 
originally published — has priority from the date of its original publication. 

Names of groups higher than genera are in the plural. Some are proper nouns; the 
remainder are adjectives used as proper nouns, agreeing in gender with the names of 
the kingdoms in which they are included; either expressing characters of the groups 
which they designate, or consisting of generic names modified by terminations signi- 
fying "resembling" or "of the group of." Plurals of generic names are not tenable 
(de Candolle, 1813) : Ericae means the species of the genus Erica; it does not mean, 
and can not be used to designate, the genus together with its allies. Names consisting 
of words other than generic names modified by terminations signifying "resembling" 
or "of the group of" are not tenable, because they are nonsense: the name Conifer- 
inae, applied by Engler to a class, is an adjective with an additional adjectival termi- 
nation superimposed. 

10 ] The Classification of Lower Organisms 

A name once applied in any principal category may not be transferred to another, 
unless it be of a form barred in the former and prescribed in the latter. The main 
clause of this statement is a consequence of the rule of priority. The exception is a 
concession to the practice of using names with uniform endings in certain categories. 

Names of groups not of principal categories do not have priority as against names 
applied in principal categories. This practice, which denies to names in subordinate 
categories the full sanction of priority, is justified by the fact that groups in these cate- 
gories are of concern only to specialists in the groups in which they occur; one is not 
in reason responsible for being aware of their names in groups outside of ones own 

Almost all families of plants have had names with the uniform ending -aceae from 
the point of time at which the category of families was distinguished from that of 
orders. Such names were applied to algae, liverworts, and mosses by Rabenhorst 
(1863) and to higher plants by Braun (in Ascherson, 1864). They are adjectives in 
the feminine, agreeing with the name of the kingdom Plantae. It is altogether expe- 
dient that names of this form be held obligatory throughout the kingdom of plants. 
A uniform termination for names of families of animals has been in use for many 
years, but these names are not equally positively sound both grammatically and by 
priority. There has been a strong tendency to apply uniform terminations to the names 
of groups of other categories. So far as concerns groups of subordinate categories — 
suborders, subfamilies, and so forth — this practice appears expedient; these groups 
being of concern only to experts in the groups in which they occur, it is as well that 
their designations be of the nature of code designations rather than names. In at- 
tempting to put this practice into effect, some zoologists have made the mistake of 
applying the same adjective in different genders to different groups; they have not 
realized that Amoebida is the same word as Amoebidae. Meanwhile, uniform termi- 
nations for names of phyla, classes, and orders, beside involving wholesale violation 
of priority, is something of an insult to the intelligence. 

The terminations of ordinal names in -ales and of family names in -aceae, currently 
in use among the Mychota, are here changed to -alea and -acea to agree with the 
neuter name of the kingdom. A change of the gender of an adjective does not create 
a new word, and the original authorities for the names will stand. Accordingly: 

The name of an order of Mychota, if based on that of a genus, must bear the termi- 
nation -alea. Names of this form are valid in no other category of this kingdom, and 
may be reapplied to orders. They have priority and authority by publication explicitly 
as orders. Such names do not supersede older ordinal names not based on names of 

The name of a family of Mychota is formed of the stem of a generic name (not 
necessarily a valid name, but never a later homonym) by adding the termination 
-acea. Names of this form are not valid in any other categor)', and may be reapplied 
to families. They have priority and authority by publication explicitly as families. 

The names of families of Protoctista, unlike those of Mychota, of plants, and of 
animals, do not have by priority prevalently a uniform termination. Many of the 
oldest were first named in -ina. Those of flagellates and myxomycetes have double 
sets of names, respectively in -aceae and -idae, in current use. It is not expedient to 
impose uniform terminations on the names of these groups, at least not in the present 
work. Accordingly: 

Each group of Protoctista is called by its oldest name of tenable form in the cor- 
rect category, barring any previously used in other principal categories, irrespective 

An Essay on Nomenclature [ 1 1 

of termination. All names which are adjectives are used in the neuter, but ascribed to 
the original authors. 

The practices described have resulted in the use of many names which will seem 
strange, producing lists which are undeniably heterogeneous. A friendly critic notes 
as an example of these things the Hst of classes, Heterokonta, Bacillariacea, Oomy- 
cetes, and Melanophycea, on page 55. It will be realized that the three among these 
names which are adjectives must be in the feminine if the groups are construed as 
Plantae, neuter if Protoctista. Taking this fact into account, these are actually the 
first names, not previously used in other principal categories, applied to these groups 
as classes. What other names could one use? Everyone will know what groups are 
intended. Would any person understand them better if new names had been created 
by applying a uniform termination to the old roots? 

Enough about nomenclature. We should begin to deal with organisms. 

Chapter III 

Kingdom I. MYCHOTA Enderlein 

Stamm Moneres Haeckel Gen. Morph. 2: xxii ( 1866), in part. 

ScHizoPHYTAE Cohn in Beitr. Biol. Pfl. 1, Heft 3: 201 (1875). 

Class ScHizoPHYTA or Protophyta McNab in Jour, of Bot. 15 : 340 ( 1877 ) ; not sec- 
tion Protophyta nor cohors Protophyta Endlicher (1836). 

Kingdoms Protophyta and Protozoa Haeckel Syst. Phylog. 1: 90 (1894), in part; 
not Protophyta Endlicher nor class Protozoa Goldfuss (1818). 

Subdivision Schizophyta Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, 
Abt. la: iii (1900). 

Division Schizophyta Wettstein Handb. Syst. Bot. 1 : 56 ( 1901 ). 

Phylum Protophyta Schaffner in Ohio Naturalist 9: 446 (1909), in part. 

Kingdom Mychota Enderlein Bakt.-Cyclog. 236 (1925). 

Kingdom Monera Copeland f. in Quart. Rev. Biol. 13: 385 (1938). 

Kingdom Anucleobionta Rothmaler in Biol. Zentralbl. 67: 248 (1948). 

Organisms without nuclei. 

The common name of Mychota in general is bacteria, but those which contain 
chlorophyll together with other pigments which make the green color impure are 
called blue-green algae. 

The cells of Mychota are always separate or physiologically independent: multi- 
cellular bodies with distinct tissues do not occur. The cells are of various shapes; most 
often they are cylindrical, being of diameters from a fraction of one micron to a few 
microns, rarely more. Except in the groups of myxobacteria and spirochaets, they 
are walled; the thickness of the walls is of the order of 0.02^ (Knasyi, 1944). The 
walls may contain cellulose, but consist chiefly of pectates, compounds of slightly 
oxidized polysaccharides with sulfate, calcium, and magnesium (Kylin, 1943). These 
compounds are readily rendered gelatinous by hydration or hydrolysis, and the cells 
are often imbedded in gelatinous layers called sheaths or capsules. 

In describing the Mychota as lacking nuclei, one commits himself to one side of a 
controversy of many years duration. Because of the greater size of the cells of the 
blue-green algae, the facts are more easily ascertained in this group than in the proper 

The cells of blue-green algae (Gardner, 1906; Swellengrebel, 1910; Haupt, 1923) 
are divided into outer and inner parts which are not sharply distinct. Pigments occur 
in a dissolved or colloidal condition in the outer part, which contains also granules 
of stored food. The granules are not carbohydrate, although a form of glycogen dis- 
tinct from that of higher organisms has been extracted (Gardner; Kylin, 1943). The 
inner part contains rods and granules, some of which stain like chromatin, while 
others ("red granules of Biitschli") are stained red by methylene blue. Cell division 
is by constriction. Olive (1904) interpreted the inner part of the cell as a nucleus 
continually in process of mitosis, and accordingly without a membrane. It is true that 
in series of disk-shaped cells one may recognize series of corresponding granules. 
Where the cells are more elongate, the rods and granules of the interior are divided 
at random. Haupt expressed the impropriety of calling any part of these cells a 

Kingdom Mychota 


Recent studies of typical bacteria by conventional microtechnical methods (Rob- 
inow, 1942, 1949; Tulasne and Vendrely, 1947) and by the electron microscope (Hil- 
lier, Mudd, and Smith, 1949) have made it possible to recognize the essential identity 
of the structure of their cells with those of the blue-green algae. The protoplast con- 
sists of outer and inner parts. The outer part, considered as a substance, may be 
called ectoplasm (Knasyi, 1930), and the inner, considered as a body, may be called 
the central body (Biitschli, 1890). The ectoplasm is very thin, occupying usually less 
than one fifth of the radius of the cell. The spiral bands which have often been seen 








•■■•;i©\ ^-> 


V ^ .?.. ■fw .^,.'^» ■; 


■4 m 

Fig. 1.- — Structure of cells of blue-green algae, a, Symploca Muscorum after 
Gardner (1906). b, Oscillatoria Princeps after Olive (1904). C, Lyngbya sp. from 
a slide prepared by Dr. P. Maheshwari, x 1,000. d, Anabacna circinnalis after 
Haupt (1923) x 2,000. 

in cells of bacteria, and which Swellengrebel ( 1906) mistook for a nucleus, are thick- 
enings of the ectoplasm. Specific stains for nucleoprotein (chromatin), as Feulgen 
or Giemsa, usually color uniformly the entire central body. If the cells are exposed to 
hydrochloric acid, a part of the nucleoprotein, containing ribonucleic acid, dissolves. 
The remainder, containing desoxyribonucleic acid, persists in the form, basically, of 
a single fairly large granule in each cell. In rod-shaped bacteria, this granule appears 
usually to divide by constriction before the cell begins to divide, and may redivide, 
so that the cell may contain two dumb-bell shaped bodies. De Lamater and Hunter 
(1951) succeeded in a partial de-staining of the dumb-bell shaped bodies and inter- 
preted them as dividing nuclei containing centrosomes and definite numbers of 
chromosomes; typical chromosomes, however, are never as small as the bodies they 
describe, and are not imbedded in bodies of nucleoprotein from which they can be 
distinguished only by the most refined technique. Enderlein (1916) observed in rod- 
shaped bacteria series of granules of which some at least are identical with the dumb- 

14 ] The Classification of Lower Organisms 

bell shaped bodies. He named these granules mychits. It might be held that the 
mychit is a chromosome, and the central body of bacteria a nucleus of a single 
chromosome, if it were not true that the blue-green algae contain comparable bodies 
of variable form and indefinite number. 

Many bacteria swim by means of flagella. The diameter of the flagella, as revealed 
by the electron microscope, is of the order of 0.02 [J.. Their positions and lengths were 
made known, before the invention of the electron microscope, by the technique of 
Loeffler (1889), which consists essentially of depositing upon them a heavy layer 
of tannic acid. By the absence or presence and arrangement of flagella, bacteria are 
classified as of four types: atrichous, without flagella; monotrichous, with one flagel- 
lum at one end; lophotrichous, with a tuft of flagella at one end; peritrichous, with 
flagella on the sides. 

Myxobacteria, spirochaets, and such blue-green algae as are sheathless filaments, 
are capable of bending movements (some spirochaets, observed with the electron 
microscope, are found also to have flagella at the ends of the cells). Spirochaets swim 
vigorously; in myxobacteria and blue-green algae, the bending movements are a mat- 
ter of slow writhing. Filaments and cells of blue-green algae are capable also of a 
moderately rapid gliding movement. The mechanism of this movement has been 
the subject of much speculation, reviewed by Burkholder ( 1934), but remains uncer- 
tain. The appearance of the movement is as though it were caused by local secretion 
of substances affecting surface tension. 

The normal reproduction of Mychota is by constriction of the cells, each into two 
equal daughter cells; whence the various names in schizo- (Greek axi^co, to split). 
Henrici (1928) studied the changes undergone by bacteria during multiplication. As 
the cells become numerous, decreasing the food supply and producing substances 
harmful to themselves, they begin to attain greater length before dividing. Subse- 
quently there is a gradual transition to enlarged and distorted forms called involution 
forms, which divide irregularly, cutting off minute fragments. These observations 
suggest the idea that the involution forms are the true normal forms of bacteria, the 
so-called normal forms being a temporary stage adapted to rapid multiplication 
under favorable conditions. 

In many rod-shaped bacteria, when conditions cease to be ideal, the protoplasts 
produce within themselves walled bodies of dehydrated protoplasm called spores 
(endospores). In general, each cell produces only one spore. No experiment has 
definitely shown how long these spores can remain alive; it is surely a matter of cen- 
turies, doubtfully of millenia. 

Lohnis and Smith (1916, 1923) observed of Azotobactcr that numbers of proto- 
plasts might escape from their walls and unite in a common mass, which they named 
the symplasm. The existence of this stage has never been confirmed by other authori- 
ties. If the symplasm exists, it is a device for achieving the effect which nucleate or- 
ganisms attain by sexual reproduction, that is, combination of the heredity of differ- 
ent lines of ancestry. 

Tliat Mychota can actually combine characters from different linos of ancestry 
was first demonstrated beyond question by Tatum and Ledcrberg (1947). They 
mixed cultures of pairs of varieties of Escherichia coli, differing in two or more 
physiological characters, and isolated from the mixtures races having characters de- 
rived from both components. Further Mork, reviewed by Ledcrberg and Tatum 
(1953), has abundantly demonstrated phenomena analogous to typical sexual 

Kingdom Myrhola 



^ V* 

••% ^ 

^ ^^ m^ W 9m w% 

#^., li.^ 

Fig. 2. — Photographs of Escherichia coli by Dr. C. F. Robinow, reproduced by 
Hillier, Mudd, and Smith (1949); left, stained to show the ectoplasm, in which 
there are thickenings which tend to be spiral; right, stained to show the large re- 
peatedly dividing granule in the central body. About x 2,000. By courtesy of Dr. 
Robinow and of the Society of .\merican Bacteriologists. 

Kingdom Mychota [17 

The metabolic systems of the Mychota are remarkably diverse. The most super- 
ficial list of physiological types would include the following: (a) anaerobic parasites 
and saprophytes; (b) facultatively aerobic parasites and saprophytes; (c) the vinegar 
bacteria, being apparently the only known organisms which, while requiring organic 
matter, are incapable of anaerobic energesis; (d) the autotrophic bacteria, the only 
organisms which maintain life by oxidation of inorganic matter; (e) organisms living 
by incomplete photosynthesis; and (f) organisms capable of typical photosynthesis. 

Geologically, the Mychota are ancient. Iron deposits and certain other formations 
believed to have been produced by them occur in Archeozoic rocks estimated as more 
than a billion years old. 

More than five thousand names have been applied to species of bacteria, but in 
the attempt to distinguish them, only about fifteen hundred are enumerated (Ber- 
gey's Manual, 6th ed., 1948). The species of blue-green algae are probably fewer 
than one thousand. 

The classification of this group is inescapably highly tentative. The morphology 
is simple and not highly varied; the physiological characters likewise appear simple, 
but are highly varied, including many which are not known in other groups. The 
antiquity of the Mychota makes it probable that many groups which appear to be- 
long together consist actually of parallel developments. The undoubted antiquity of 
the apparent main groups would lead one to place them in the category of divisions 
or phyla; but it is not expedient to make many divisions of a group of 2500 species: 
this would produce too many divisions of a single class or classes of a single order. 
The kingdom is accordingly treated as a single phylum, and its main divisions as 

Phylum ARCHEZOA Haeckel 

yhylB. Archephyta and Archezoa Haeckel Syst. Phylog. 1:90 (1894); not Phylum 

Archephyta Haeckel (1866). 
Phylum Myxophyceae Bessey in Univ. Nebraska Studies 7: 279 (1907). 
Phyla Dimychota and Monomychota Enderlein Bakt.-Cyclog. 236 (1925). 
Bacteriophyta and Cyanophyta Steinecke (1931). 
Stamme Cyanophyta and Schizomycophyta Pascher in Beih. bot. Centralbl. 48, 

Abt. 2: 330 (1931). 
Divisions Cyanophyta and Schizomycetae Stanier and van Niel in Jour. Bact. 42: 

464 (1941). 
Characters of the kingdom. 

Archezoa is Haeckel's name, at the point cited, for the bacteria. The name had 
been applied othervv^ise by Perty (1852), but not in a principal category. It will not 
be considered inappropriate, if it be remembered that the meaning of zoe is as much 
life as animal. 

The conventional division of the group into two classes, bacteria and blue-green 
algae, is not perfectly natural. All of the recognized blue-green algae belong together; 
but the recognized bacteria are a wide miscellany, some of them belonging with the 
blue-green algae. Here three classes are recognized. 
1. Cells without internal pigment, heterotrophic 
or living by chemosynthesis; not usually pro- 
ducing filaments with prominent sheaths. 

18 ] The Classification of Lower Organisms 

2. Cells with firm walls, non-motile or 

motile by means of flagella Class 1. Schizophyta. 

2. Cells with thin walls or none, motile by 
means of changes of shape, also some- 
times by flagella Class 2. Myxoschizomycetes. 

1. Cells mostly with internal pigment, living by 
photosynthesis or chemosynthesis, exception- 
ally heterotrophic; often producing filaments 
with prominent sheaths Class 3. Archiplastidea. 

Class 1. SCHIZOPHYTA (Cohn) McNab 

Schizomycetes Nageli ex Caspary in Bot. Zeit. 15: 760 (1857). 

Class Schizophyta or Protophyta McNab in Jour, of Bot. 15: 340 (1877). 

Class Schizomycetes Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt, 1: 

33 (1879). 
Class Schizomycetae SchafTncr in Ohio Naturalist 9: 447 (1909). 
Classes Holocyclomor pha and Hemicyclomorpha Enderlein Bakt.-Cyclog. 236 

Dependent or chemosynthetic Mychota, with walled cells, without photosynthetic 
pigments and not producing sheathed filaments. 

This class includes as orders the typical bacteria and two minor groups. 
1. Cells solitary or loosely gathered into clusters 

or filaments, spherical, rod-shaped, or spiral, 

not differentiated along the axis Order 1. Schizosporea. 

1. Consisting of branched filaments not divided 

into cells Order 2. Actinomycetalea. 

1. Cells attached by stalks, the attached and 

free ends differentiated Order 3. Caulobacterialea. 

Order 1. Schizosporea [Schizosporeae] Cohn in Hedwigia 11: 17 (1872). 
Order Schizomycetes (Nageli) McNab in Jour, of Bot. 15: 340 (1877). 
Order Eubacteria Schroter 1886. 

Order Haplobacteriacei Fischer in Jahrb. wiss. Bot. 27: 139 (1895). 
Orders Cephalotrichinae and Peritrichinae Orla-Jensen in Centralbl. Bkt. Abt. 

2,22: 334,344 (1909). 
Order Eubacteriales Buchanan in Jour. Bact. 2: 162 (1917). 
Mychota whose cells in the typical condition are without internal pigment, walled, 
of the form of rods, spheres, or spirals, not differentiated along the axis. As this is a 
numerous group, likely with advancing knowledge to require division, it will be well 
to provide it with a nomenclatural standard, and to suggest as such Cohn's principal 
discovery among bacteria, namely Bacillus sublilis. 

These are the typical bacteria. As originally described by Leeuwcnhoeck (1677), 
they were taken to be a few kinds of "animacules" distinguished only by extremely 
small size. Only after many years were they shown to be numerous and varied, and 
highly important as causes of diseases and of other natural phenomena. 

The natural classification of the typical bacteria has been hard to discern. The 
characters by which groups can be distinguished include forms of cells and of clusters 
of cells; absence or presence and arrangement of flagella; non-formation or formation 

Kingdom My c hot a [ 19 

of endospores; metabolic products; and the peculiar character called Gram reaction. 
The method of staining invented by Gram, 1884, consists of staining successively 
with gentian violet and iodine. It gives an intense blue-black color. From some bac- 
teria, this color is washed out by alcohol; others retain it; the former are said to be 
Gram negative, the latter Gram positive. In practice one applies successively gentian 
violet, iodine, alcohol, and safranine, the last being a red dye whose function is to 
make the Gram negative bacteria visible. The substance stained by gentian violet 
plus iodine is believed to be lipoid, such as occurs in all cells. The Gram positive 
quality is believed to consist in a relatively low isoelectric point, a capacity, that is, 
to combine with anions in a relatively acid medium. This quality lies in the ectoplasm 
of the cells and disappears in aging cultures. 

The classification given in Bergey's Manual (1923, 1925, 1930, 1934, 1939, 1948) 
is accepted (at least among Americans) as standard. The following system of thirteen 
families is a moderate rearrangement of the Bergeyan system, with certain ideas or 
names from Enderlein (1917, 1925), Buchanan ( 1925), Pribram (1929) and Stanier 
andvanNiel (1941). 

1. Gram positive, with exceptions many of 
which are intracellular parasites; atrichous or 

2. Spheres dividing in more planes than 

3. Gram positive Family 1. Micrococcacea. 

3. Gram negative; intracellular patho- 
gens in animals Family 2. Neisseriacea. 

2. Rods, or spheres dividing in one plane. 
3. Not producing endospores. 
4. Atrichous. 

5. Not intracellular parasites. . Family 3. Corynebacteriacea. 

5. Intracellular parasites Family 4. Rickettsiacea. 

4. Peritrichous Family 5. Kurthiacea . 

3. Producing endospores Family 6. Bacillacea. 

1. Gram negative. 

2. Atrichous or peritrichous, requiring com- 
paratively complicated organic food. 
3. Not plant pathogens. 
4. Not fixing nitrogen. 

5. Capable of growth on or- 
dinary media Family 7. Achromobacteriacea. 

5. Requiring special media; 

minute atrichous pathogens. Family 8. Pasteurellacea. 

4. Fixing nitrogen Family 10. Azotobacteriacea. 

3. Plant pathogens Family 9. Rhizobiacea. 

2. Atrichous, monotrichous, or lophotrich- 
ous; the atrichous representatives, and 
many others, can survive with organic 
foods simpler than carbohydrates, or 
with none. 

3. Mostly requiring at least carbo- 
hydrates Family 11. Spirillacea. 

20 ] The Classification of Lower Organisms 

3. Not requiring carbohydrates. 

4. Oxidizing alcohol to acetic 

acid, and acetic acid to CO2 

and H2O Family 12. Acetobacteriacea. 

4. Not as above; many examples 

strictly autotrophic Family 13. Nitrobagteriacea. 

Family 1. Micrococcacea [Micrococcaceae] Pribram in Jour. Bact. 18: 370, 385 
(1929). Family Coccaceae Zopf 1884; but the genus Coccus is a scale insect. Gram 
positive spheres producing packets or irregular masses. Micrococcus, saprophytic or 
parasitic, producing irregular masses of cells; the pathogenic species have been treated 
as a separate genus Staphylococcus. Sarcina, saprophytic or commensal spheres pro- 
ducing packets. 

Family 2. Neisseriacea [Neisseriaceae] Prevot ex Bergey et al. Manual ed. 5 : 278 
(1938). Family Neisseriacees Prevot in Ann. Sci. Nat. Bot. ser. 10, 15: 119 (1933). 
Obligate parasites, the Gram negative spherical cells occurring chiefly in pairs within 
leucocytes in the lesions of disease. Neisseria gonorrhoeae, the gonococcus; A^. ]Veich- 
selbaumii Trevisan {N. intracellularis, N. meningitidis, Auctt.), the meningococcus. 

Family 3. Corynebacteriacea [Corynebacteriaceae] Lehmann and Neumann 1907. 
Family Corynebacteriidae Enderlein in Sitzber. Gess. naturf. Freunde Berlin (1917) : 
314. Family Lactobacillaceae Winslow et al. in Jour. Bact. 2: 561 (1917). Family 
Lactobacteriaceae Orla-Jensen 1921. Family Leptotrichaceae Pribram in Jour. Bact. 
18: 372 (1929), not family Leptotrichacei Schroter 1886. Gram positive rods, or 
spheres dividing in one plane and producing chains, non-motile. 

Streptococcus, spheres in chains; saprophytes in milk, involved in the making of 
butter and cheese; and commensals and serious pathogens causing, for example, 
abscesses, septicemia, erysipelas, and pneumonia. 

Diplococcus, spheres usually in pairs, encapsulated. D. pneumoniae occurs in many 
immunologically distinct races which are the usual causes of pneumonia. 

Lactobacillus, rods, microaerophilic, producing lactic acid. In milk, involved in 
the making of butter and cheese; in the oral cavity, being the usual agent of dental 
caries (Rosebury, Linton, and Buchbinder, 1929); common in sewage. 

Leptotrichia, rods which become exceptionally long before dividing. Oral cavity 
of man and beasts. 

Corynebacterium, rods, becoming club-shaped, staining in a banded pattern. The 
type species is the agent of diphtheria, C. diphthcriae; the genus includes also many 
harmless commensals important only as making diagnosis difficult. The cells divide 
in an exceptional fashion, by breaking violently from one side to the other near one 
end; the cut-off end swings around beside the main body and proceeds to grow. 
Repeated division in this manner produces clusters of parallel cells (Park, \V'iliiams, 
and Krumweide, 1924). 

Family 4. Rickettsiacea [Rickettsiaceae] Pinkerton 1936. Families Bartonellaceae 
Gieszszykiewicz 1939 and Chlamydozoaceae Moshkovsky 1945. Minute obligate intra- 
cellular parasites of varied form, commonly Gram negative but with Gram positive 

There have been many observations of bodies of the characters stated, but a satis- 
factory classification of them is not yet possible. Howard Taylor Ricketts showed 
that Rocky Mountain spotted fever is transmitted by the tick Dcrmocentor, and 
observed, in the cells of diseased tissues, minute irregularly staining bodies; in 1910, 

Kingdo7n Mychota [21 

in the course of further studies of the disease, he contracted it and died. Stanislas 
Prowazek, called into the Austrian military medical service in 1914, began to study 
typhus, which is transmitted by lice; observed similar intracellular bodies; contracted 
typhus, and died in February, 1915 (Hartmann, 1915). The cause of Rocky Mountain 
spotted fever is Rickettsia Rickettsii, and that of typhus. is R. Prowazekii. Several 
other species are known. By serological methods, Anigstein (1927) showed that 
R. Melophagi is closely related to Corynebacterium. 

In cases of the disease of the west slope of the Andes called verruga peruana, 
Oroya Fever, or Carrion's disease, there occur intracellular bodies named Bartonella 
bacillijormis. Noguchi and others (192H) completed the demonstration that the 
disease is transmitted by biting flies of the genus Phlebotoyniis. Good authority has 
construed Bartonella as a sporozoan. 

Students of flagellates, Sarkodina, and Infusoria have occasionally observed in 
the cytoplasm or nuclei of these organisms minute bodies multiplying to form consid- 
erable masses. These parasites have generally been construed as chytrids, but have 
little in common with proper chytrids. The genus Caryococcus Dangeard includes at 
least a part of them. 

Family 5. Kurthiacea, fam. nov. Gram positive peritrichous rods, not producing 
endospores. Kurthia, harmless; Listeria Pirie ex Murray in Bergey's Manual 6th ed. 
408 (1948), pathogenic in sheep and man. 

Family 6. BaciUacea [Bacillacei] Fischer in Jahrb. wiss. Bot. 27: 139 (1895). 
The spore-forming rods, always Gram positive, mostly peritrichous, very numerous in 
species, common, and important. 

Bacillus Cohn 1872, is one of the oldest generic names of rod-shaped bacteria 
which can be definitely applied: it can be definitely applied because the type species 

B. subtilis was so described as to be recognizable. The genus has been used to include 
rods in general or at random. Defined as aerobic spore-formers, as proposed by 
Buchanan, 1917, it is a thoroughly natural group. As treated in the fifth edition of 
Bergey's Manual, it included nearly 150 duly distinguished species; in the sixth 
edition, this number is cut to thirty-three. The great majority are saprophytic. Ex- 
ceptions, important pathogens, are B. anthracis; and B. alvei and other species causing 
foulbrood of bees. 

The anaerobic spore-formers constitute the genus Clostridium. The type species 
wa? discovered and named three times in different connections. As an anaerobe 
involved in the fermentations which give butter its flavor, it is C. butyricum Prazmow- 
ski. As an organisms whose cells contain granules staining like starch, it is Bacillus 
Amylobacter van Tieghem. It has the property of fixing nitrogen; discovered in this 
capacity by Winogradsky (1902) it was named C. Pastorianum. The species of 
Clostridium, as of Bacillus, are numerous. They are primarily saprophytic, but many 
species produce powerful toxins and are serious pathogens. Examples are C. tetani; 

C. botulinum; and C. septicum and a whole roll of other species, causing various 
forms of gangrene, occasion for the study and distinction of which was found during 
World War I. 

Family 7. Achromobacteriacea [Achromobacteriaceae] Breed 1945. Family Bac- 
teriaceae McNab in Jour, of Bot. 15: 340 (1877), based on a generic name which 
must be abandoned as a nomen conjusum. Family Enterobacteriaceae Rahn 1937, not 
based on a generic name. Gram negative rods which lack the dictinctive characters 
of the families subsequently to be treated. 

22 ] The Classification of Lower Organisms 

The nine genera listed first occur normally in animals, mostly in the gut and 
mostly as commensals; exceptions are important pathogens. Most of them produce 
acid, and many of them produce gas, from sugar. These genera are the traditional 
colon-typhoid-dysentery group. 

Escherichia coli, the colon bacillus, and Aerohacter aerogenes, the gas bacillus, are 
common commensals which produce acid and gas from dextrose and lactose. The 
standard method of testing waters for contamination is essentially a test for the 
presence of these organisms. 

Klebsiella also produces acid and gas from sugars. It inhabits the respiratory 
tract. The cells are heavily capsulated and non-motile. The type species K. pneumo- 
niae is an important pathogen, the pneumobacillus of Friedlander. 

Proteus vulgaris (this is at least the third genus to bear the name Proteus, but the 
first in this kingdom) produces acid and gas from dextrose but not lactose, and 
liquefies gelatine. It is usually isolated from spoiled meat. 

Salmonella is distinguished from Proteus by non-liquefaction of gelatine. Many 
of its species are harmless commensals; others cause paratyphoid fevers. Immunologi- 
cal study of cultures of Salmonella from cases of disease and from waters have re- 
sulted in the distinction of fully 150 races, mostly unnamed and identifiable only by 
immunological reactions. Eberthella includes motile rods producing acid but not 
gas from sugars, and belonging to the same immunological system as the various 
races of Salmonella. Eberthella typhi causes typhoid fever. 

Shigella is distinguished from Eberthella by non-motility. The Shiga bacillus, 
S. dystenteriae, is the cause of dystentery. 

Bacteroides is a numerous group of acid-producing gut bacteria, motile or non- 
motile, generally harmless.^ distinguished from the foregoing as strictly anaerobic. 

Alcaligenes fecalis, an apparently harmless organism isolated from intestinal con- 
tents, does not produce acid from sugars; grown in milk, it produces an alkaline 

Numerous races of bacteria which have been isolated from soil and are capable 
of attacking cellulose are assigned to the genus Cellulomonas. Bacteria which produce 
an extracellular red pigment are Serratia (one of the oldest generic names for bac- 
teria); those which produce yellow pigment are Flavobacterium; those which produce 
blue, black, or violet growths are Chromobacterium. Cultures which lack the distinc- 
tive characters of all of the above named genera (most such cultures have been 
isolated from water) are called Achromohacter. 

Family 8. Pasteurellacea nom. nov. Family Parvobacteriaceae Rahn; there is no 
corresponding generic name. Minute non-motile Gram negative rods, pathogenic, 
requiring special media for cultivation. Pasteurclla avicida is the cause of chicken 
cholera, upon which Pasteur made important studies. Of greater direct importance 
to man is Pasteurella pestis, the cause of plague. Hemophilus includes the agents 
of whooping cough, soft chancre, and conjunctivitis. Brucella includes the organisms 
which cause Malta fever, undulant fever. Bang's disease, contagious abortion. Pfeif- 
ferella mallei is the cause of glanders. 

Family 9. Rhizobiacea [Rhizobiaceae] Conn in Jour. Bact. 36: 321 (1938). Gram 
negative rods, atrichous or peritrichous, parasites on plants. Cultured in the presence 
of sugars, these organisms produce acid; they are evident allies of the colon group. 

Erwinia commemorates Erwin F. Smith, the discoverer of many bacteria pathogenic 
to plants. Typical species cause blights, wilts, or dry necroses. The discovery by 
Burrill, 1882, of Erwinia amylovora, the cause of the fire blight of pears, should 

Kingdom Mychota [ 23 

have prevented the formulation of a theory, once entertained, that all bacteria 
require neutral media, and are accordingly incapable of causing diseases of plants. 
The species of Pectobacterium, as P. carotovorum, cause rots. Those of Agro- 
bacterium cause galls; A. tumefaciens causes crown gall of many plants. 

Rhizobium includes the species which produce little galls ("nodules") on the 
roots of plants and which benefit their hosts by fixing nitrogen. The best known 
hosts of Rhizobium are plants of the family Leguminosae; the relationship between 
Leguminosae and Rhizobium is a classic example of symbiosis. There are several or 
many species of Rhizobium, scarcely distinguishable morphologically, but living on 
different groups of legumes. The race which was first recognized and isolated, R. 
Leguminosarum Frank 1890 [Schinzia Leguminosarum Frank 1879; Bacillus Radicic- 
ola Beijerinck 1888) is that which attacks plants of the pea tribe. Bewley and Hutch- 
inson (1920) accounted for the variety of forms which Rhizobium can assume. In 
the roots of plants it occurs as involution forms. In culture, it is a peritrichous rod, 
but the flagella are often reduced to one, and it has been confused with the mono- 
trichous bacteria (Conn and Wolfe, 1938). 

Family 10. Azotobacteriacea [Azotobacteriaceae] Bergey, Breed, and Murray in 
Bergey's Manual 5th ed., preprint, v and 71 (1938). These are the organisms which 
were originally isolated by Beijerinck (1901) by inoculating with garden soil shallow 
layers of a nitrogen-free nutrient solution containing mannite. The commonest species, 
Azotobacter Chroococcum, is usually seen as ellipsoid cells, as much as \\x thick and 
7[J. long, solitary, with peritrichous flagella, or forming non-motile clusters imbedded 
in a heavy capsule. Beijerinck observed the occurrence of globular involution forms 
as much as 15^ in diameter. Lohnis and Smith (1916) made a thorough study of 
variations in form, and reported a remarkable variety of other stages, including the 

The Pasteurellacea and Rhizobiacea are apparently reasonably close allies of 
the Achromobacteriacea. The Azotobacteriacea stand somewhat apart. The remain- 
ing families of the present order are more definitely distinct, being marked by mono- 
trichous or lophotrichous flagella. 

Family 11. Spirillacea [Spirillaceae] Migula 1894. Family Pseudomonadaceae 
Winslow et al. in Jour. Bact. 2: 555 (1917). Rods and spirals, Gram negative, mono- 
trichous or lophotrichous; not producing much acetic acid, and mostly heterotrophic. 

Pseudomonas is a numerous genus of rods which may or may not produce a fluores- 
cent pigment soluble in water; they do not produce a yellow pigment which is in- 
soluble in water. The original species, P. aeruginosa, was isolated from pus, in which 
it produces a blue-green discoloration; it is by itself weakly if at all pathogenic. 
Other species have been isolated from fresh and salt waters and brines; the bacteria 
which produce phosphorescence on salt fish are of this genus. Many further species 
arc: pathogenic to plants, producing chiefly leaf spots. 

Phytomonas Bergey et al. 1923 {Xanthomonas Dowson 1948) includes numerous 
plant pathogens which in culture produce an insoluble yellow pigment; among them 
are the causes of cabbage rot, walnut blight, and leaf spots on many plants. 

Pacinia Trevisan 1885 includes monotrichous curved rods. The type species P. 
cholerae-asiaticae is the cause of Asiatic cholera. Among numerous other species 
the majority are harmless saprophytes in waters. Recent authorities have treated the 
cholera organism as the type of the genus Vibrio Miiller (1773); their action is an in- 
tolerable falsification of the usage of a full century preceding the discovery of the 
cholera organism. 

24 ] The Classification of Lower Organisms 

Spirillum includes the typical spirals, lophotrichous, a small number of species of 
harmless saprophytes in foul waters. 

Thiospira includes large lophotrichous spirals, colorless, containing granules of 
sulfur. They are believed to live by chemosynthesis. 

Family 12. Acetobacteriacea [Acetobacteriaceae] Bergey, Breed, and Murray 
1938. As gross objects, growths of Acetobacter aceti Beijerinck have been known since 
prehistoric times. With included yeasts they constitute mother of vinegar (the old 
names Mycoderma mesentericum Persoon, Ulvina aceti Kiitzing, and Umbina aceti 
Nageli designated the combination of bacteria and yeasts, and it seems proper to 
reject them). Free-swimming cells with polar flagella have been observed; ordinarily 
the cells appear as rods in chains, heavily encapsulated, or as involution forms. 
The organic food required by Acetobacter is alternatively alcohol, which is oxidized 
to acetic acid, or acetic acid, which is oxidized to carbon dioxide and water. These 
processes are strictly aerobic: to make vinegar, one exposes wine to air; to preserve 
it, one seals the vessels. 

Family 13. Nitrobacteriacea [Nitrobacteriaceae] Buchanan in Jour. Bact. 2: 349 
(1917). Organisms oxidizing the simplest organic compounds; or facultatively capa- 
ble of chemosynthesis; or living strictly by chemosynthesis and strictly aerobic: mostly 
Gram negative monotrichous or atrichous rods. 

Methanomonas is capable of oxidizing methane; Carboxidomonas of oxidizing 
carbon monoxide; Hydrogenomonas, of oxidizing elemental hydrogen. Thiobacillus 
includes organisms which oxidize hydrogen sulfide or elemental sulfur. 

Winogradsky had discovered chemosynthesis in the course of studies of Beggiatoa 
and other sulfur-oxidizing organisms before he undertook to isolate bacteria which 
cause nitrification, that is, the natural production of nitrates in soil and waters. 
He achieved success (1890) by inoculating, with soil or sewage, media which con- 
tained salts of ammonia but no food; he saw the nitrifying organisms first as minute 
motile rods which he named Nitromonas. Further study and the use of solid media 
showed that nitrification takes place in two stages and is the work of several kinds of 
organisms. Winogradsky distinguished Nitrosomonas europaea and N. javaneyisis, 
monotrichous rods from different regions as indicated, oxidizing ammonia to nitrites; 
Nitrosococcus, non-motile spheres from South Amerca, effecting the same oxidation 
as Nitrosomonas; and Nitrobacter, non-motile rods oxidizing nitrites to nitrates. 
Subsequent authors have validated Winogradsky's names by creating the combina- 
tions Nitrosococcus nitrosus and Nitrobacter VVinogradskyi. Subsequently, Winograd- 
sky discovered yet other bacteria capable of the same oxidations. 

The presence of nitrifying bacteria is necessary for the normal growth of most 
crops. So active are the nitrifying bacteria that no more than traces of ammonia and 
nitrites are found in normal soils, and so avidly do plants absorb nitrates that these 
accumulate only in fallow fields. 

Order 2. Actinomycetalea [Actinomycetales] Buchanan in Jour. Bact. 2: 162 


Organisms which consist typically of slender filaments not divided into cells, 
but which are capable of producing conidia, that is, minute spherical or elongate 
bodies cut off by constriction from the ends of the filaments, or of breaking up into 
cells of the form of regular or irregular rods. Non-motile; Gram positive or Gram 
negative; often of the staining character called acid fast. 

The order may be treated as a single family. 

Kingdom Mychota [ 25 

Family Mycobacteriacea [Mycobacteriaceae] Chester 1907. Family Actinomyce- 
taceae Buchanan in Jour. Bact. 3: 403 (1918). Family Streptomycetaceae Waksman 
and Henrici 1943. Characters of the order. Three genera require discussion. 

Streptomyces Waksman and Henrici 1943. The original name of this genus is 
Streptothrix Cohn (1875); there is an older genus Streptothrix among plants, and 
the numerous species of the present genus have generally been included in Actino- 
myces. Cultures are readily isolated from air or soil. They appear as slowly growing 
colonies which may at first be of various colors and have shiny surfaces. Their texture 
is tough; a blunt needle will more often tear a colony from the medium than pene- 
trate it. As the colonies grow, they become truncate; the exposed surfaces become 
white and powdery; pigments, black, brown, red, or yellow, in various races, are 
produced, and discolor the medium. The toughness of the colonies is a consequence 
of their structure, of myriad crooked branching filaments about 1|J. in diameter, 
without joints; the white and powdery surface is produced by myriad conidia released 
in basipetal succession. The cultures are of an odor which may be described as that 
of earth under the first rain after drouth: undoubtedly, this familiar odor is that of 
Streptomyces in the soil. Drechsler (1919), from careful study of several species of 
Streptomyces, concluded that they are fungi; their filaments are, however, much 
finer than those of fungi, and no definite nuclei have been seen. 

Certain species of Streptomyces cause a scabbiness of potatoes. Except for this, the 
genus was for a long time regarded as quite unimportant. When the capacity of the 
fungus Penicillium notatum to inhibit the growth of bacteria had been observed, 
and had led to the discovery of the drug penicillin, Waksman, the leading authority 
on the classification of Actinomycetalea, sought comparable drugs produced by 
Streptomyces, and had the great success of discovering streptomycin. 

Actinomyces Bovis Harz 1877 is one of several species of the same general nature 
as Streptothrix which are pathogenic to animals. It causes lumpy jaw of cattle. 

Mycobacterium Lehmann and Neumann 1896 is typified by M. tuberculosis, the 
agent of one of the most important diseases of man, supposed originally to have 
attacked cattle, and to have spread around the world with European cattle. It is a 
chronic disease, destroying the tissues slowly and producing a nugatory sort of im- 
munity which makes it possible to test for the disease, but does not check it. The 
cells are recognized in sputum and in diseased tissues by the acid fast reaction: the 
dye carbol fuchsin must be applied hot in order to color them; once it has done so, 
it does not wash out in acid alcohol. It is cultivated with difficulty. The growth is 
dry, powdery, wrinkled, with an odor described as sickening-sweet. It consists of 
branching filaments which break up readily into rod-shaped or irregular fragments. 

Lesions of leprosy contain acid fast organisms named Mycobacterium leprae. Gay 
(1935) has discussed the results of attempts to cultivate this species. They have 
yielded either "diphtheroid" cells or a "streptothrix." He concludes that most of 
the reports are of the same organism reacting variously to various conditions. 

Order 3. Caulobacterialea [Caulobacteriales] Henrici and Johnson in Jour. Bact. 
29: 4 (1935). 

Aquatic bacteria, the cells of most examples secreting gelatinous matter in such a 
manner as to produce stalks. Henrici and Johnson provided a system of four families, 
five genera, and nine species. Stanier and van Niel (1941) rejected the group as 
artificial, placing some of the genera among Eubacteria and leaving others unplaced. 
The order may be maintained for the accommodation of the latter and divided into 
two families. 


The Classification of Lower Organisms 


Fig. 3 — a-e, Caulobacterialea after Henrici and Johnson (1935) x 2,000: a, 
Nevskia sp.; b, Caulobacter vibrioides; c, Caulobacter sp.; d, Pasteuria sp.; e, Blasto- 
caulis sp. f. Various stages of Cytophaga Hutchinsonii [Spirochaeta cytophaga) after 
Hutchinson and Clayton ( 1919). g-k, Myxobactralea after Thaxtcr (1892), the cells 
X 1,000, in the fruits x 200. g, h, Cells and fruit of Chondromyccs crocatus; i, fruit 
of C. aurantiacus; j,k, vegetative cells and spores, and fruit, of Myxococcus coralloi 
des. I, m, Dividing cells of Cristispira Veneris after Dobell (1911) x 2,000. 

Kingdom Mychota [ 27 

Family 1. Leptotrichacea [Leptotrichacei] Schroter 1886. The cells not elongated 
in the direction of the axis of the stalk. 

Didymohelix ferruginea (Ehrenberg) Griffith (first named, and usually listed, 
under Gallionella, which is a misspelling of the name of a genus of diatoms) occurs 
in waters containing iron. Older authors described it as consisting of paired filaments, 
less than 1^ in diameter, colored bright yellow with imbedded iron oxide, and coiled 
about each other. In fact, the supposed paired filaments are the margins of a single 
twisted band, which is not itself an organism but the stalk secreted by a terminal 
cell. Spirophyllum Ellis is either the same species or a closely related larger one. 

Leptothrix Kiitzing Phyc. Gen. 198 ( 1843) was inadequately described; the species 
which was first named, and which is accepted as the type, was L. ochracea. It is be- 
lieved that this name properly designates the masses of ochraceous matter seen in 
iron springs. Under the microscope, this matter is seen to consist of fine yellow 
filaments, straight and unbranched. Ellis (1916) described them as consisting of a 
cylinder of protoplasm, not divided into cells, enclosed in a sheath. Almost surely, 
these structures, generally recognized as of the same nature as Didymohelix, are like- 
wise stalks secreted by minute terminal cells. 

Siderocapsa Molisch and Sideromonas Cholodny, described as minute spheres or 
rods imbedded in capsules colored by ferric oxide and attached to plants in waters 
containing iron, are perhaps to be interpreted as stalkless members of the present 

Nevskia Famintzin, forming minute gelatinous colonies floating on water, does not 
accumulate iron. 

Family 2. Caulobacteriacea [Caulobacteriaceae] Henrici and Johnson 1. c. (1935). 
The cells elongated in the direction of the long axes of the stalks. Caulobacter, Pas- 
teuria, and Blastocaulis, colorless saprophytes in waters or parasites in aquatic 


Class Myxoschizomycetae Schaffner in Ohio Naturalist 9: 447 (1909). 

Class Polyyangidae Jahn Beitr. bot. Protistol. 1: 65 (1924). 

Class Spirochaetae Stanier and van Niel in Jour. Bact. 42 : 459 ( 1941 ) . 

Parasitic or saprophytic Mychota, the elongate cells with thin walls or none, 
capable of bending movements and sluggishly or actively motile. In many examples 
there is a resting stage: the cell contracts generally, so as to diminish the surface, 
and deposits a definite wall. The structure so produced is a spore of the type called 
an arthrospore or chlamydospore. 

The two orders Myxobactralea and Spirochaetalea have not previously been 
combined to form a separate class. A certain species which Hutchinson and Clayton 
(1919) described as a spirochaet, Spirochaeta cytophaga, has subsequently been 
found to be a myxobacterium. The hint of relationship thus conveyed is confirmed 
by the whole character of both groups, as may be seen from the discussions of them 
by Stanier and van Niel (1941) and Knasyi (1944). 

Order 1. Myxobactralea [Myxobactrales] Clements Gen. Fung. 8 (1909). 

Order Myxobacteriaceae Thaxter in Bot. Gaz. 17: 389 (1892). 

Order Myxobacteriales Buchanan in Jour. Bact. 2: 163 (1917). 
The cells not definitely of spiral form, sluggishly motile. In typical examples, the 

28 ] The Classification of Lower Organisms 

cells occur in swarms imbedded in slime; the entire mass moves concertedly, and is 
eventually converted into macroscopically visible fruiting bodies. 

The group was first recognized by Thaxter. He took note that the fruiting bodies 
of Chondromyces had already been described by Berkeley and Curtis as those of a 
gasteromycete, and learned subsequently that Polyangium Link, also described as 
of the puffball group, is an older name for his Myxobacter. The swarms of cells live 
in air on damp substrata (commonly the feces of various kinds of animals), moving 
across them and digesting and absorbing food as they proceed. Labratory culture is 
fairly easy. As a reaction, apparently, to exhaustion of the available food, the cells 
change into chlamydospores; the masses of spores held together by dried slime are 
called cysts. These may be borne on simple or branched stalks built up from the 
slime as a preliminary to the formation of the cysts and spores. The group is of 
essentially no economic importance. 

The accepted classification is that of Jahn (1924); to the four families which he 
recognized, one more has been prefixed for the accommodation of the genus 

family 1. Cytophagacea [Cytophagacae] Stanier 1940. The chlamydospores 
formed sporadically by individual cells, not in cysts. Cytophaga Hutchinsonii Wino- 
gradsky [Spirochaeta cytophaga Hutchinson and Clayton) is one of several species 
discovered as active fermenters of cellulose. The slenderly spindle-shaped cells are 
sluggishly motile, and produce ellipsoid chlamydospores resembling yeasts. 

Family 2. Archangiacea [Archangiacae] Jahn op. cit. 66. Spores elongate in irregu- 
larly extensive masses, not in cysts. Archangium, Stelangium. 

Family 3. Sorangiacea [Sorangiaceae] Jahn op. cit. 73. Spores elongate, the 
cysts angular, in masses, not stalked. Sorangium. 

Family 4. Myxobacteriacea [Myxobacteriaceae] (Thaxter) E. F. Smith 1905. 
Family Polyangiaceac Jahn op. cit. 75. Spores elongate, in distinct rounded cysts, 
clustered or solitary, sessile or borne on simple or branched stalks. Polyangium 
Link 1795 [Myxobacter Thaxter 1892), Stelangium, Melitangium, Podangium, 

Family 5. Myxococcacea [Myxococcaceae] Jahn op. cit. 83. Spores spherical; 
cysts indefinite or definite. Myxococcus, Chondrococcus, Angiococcus. 

Order 2. Spirochaetalea [Spirochaetales] Buchanan in Jour. Bact. 2: 163 (1917). 

Cells solitary, spiral in shape, actively motile. 

The first known species of this group was Spirochaeta plicatilis, observed in foul 
waters by Ehrenberg (1838). The next was the species now known as Borrelia recur- 
rentis (Lebert) Bergey et al., observed in the blood of relapsing fever patients by 
Obermeier, 1873. 

During the last years of the nineteenth century, many attempts to identify the 
agent of syphilis by standard bacteriological methods were unsuccessful. The German 
government directed Schaudinn and Hoff'mann to continue this work. Fritz Schau- 
dinn, 1871-1906 (Stokes, 1931), had attained distinction as a student of pathogenic 
protozoa. Within a few weeks, by the microscopic examination of lesions, he attained 
success where the bacteriologists had failed, and discovered Treponema pallidum 
(Schaudinn and Hoffmann, 1905). 

Spirochaets were first cultivated by Noguchi; few others have been successful in 
this difficult practice. It requires a medium of aseptic, not sterilized, animal ma- 
terial, under more or less anaerobic conditions. Each species requires its peculiar 
variant of the conditions, to which it is quite sensitive. 

Kingdom Mychota [ 29 

Spirochaeta plicatilis and other saprophytic species, together with certain species 
parasitic in mollusks, are fairly large. The species which are parasitic or commensal 
in other animals may be extremely small. It is chiefly by study of the larger species 
that the structure is known. The internal structure is septate. Dobell (1911) found 
in Cristispira, at the margin of each septum, a whorl of granules staining like chroma- 
tin, and interpreted these granules collectively as a nucleus. Noguchi (in Jordan and 
Falk, 1928) saw in the interior of the smaller species no chambered structure, but a 
lengthwise rod. This has been interpreted as a nucleus, as a locomotor or skeletal 
structure, or as an artifact. The electron microscope has shown actual flagella at the 
ends of cells of Treponema pallidum. Reproduction is normally by transverse divi- 
sion into two. During division, the daughter cells may coil about one another, giving 
a false appearance of lengthwise division. Gross (1913) observed that Cristispira is 
capable of breaking up into cylindrical Stdhchen corresponding to the chambers. 

The discovery of Treponema by an eminent protozoologist; the character of 
spirochaetal diseases, several of which are spread by biting insects, and produce only 
that nugatory immunity which makes diagnosis possible but does not check the 
disease; and the supposed lengthwise division of the cells; led to the hypothesis that 
the spirochaets are protozoa. Dobell was surely correct in dismissing this hypothesis, 
insisting that the spirochaets are neither protozoa nor typical bacteria, but a group 
sui generis. 

The larger and smaller spirochaets are reasonably treated as separate families. 

Family 1. Spirochaetacea [Spirochaetaceae] Swellengrebel 1907. The cells com- 
paratively large, 80-500(1 long. Spirochaeta, Saprospira, Cristispira. 

Family 2. Treponematacea [Treponemataceae] Robinson in Bergey Man. 6th ed. 
(1948). Family Treponemidae Schaudinn 1905. The cells 4-15^ long. 

Treponema Schaudinn. The cells comparatively loosely coiled. T. pallidum, the 
agent of syphillis. T. pertenue, the agent of yaws. T. macrodentium and T. micro- 
dentium, harmless commensals in the mouth. 

Borrelia Swellengrebel is doubtfully distinct from the foregoing; Noguchi reduced 
it. B. recurrentis and other species cause relapsing fevers. B. Vincenti causes Vincent's 
angina (trench mouth). The fusiform cells always found associated with it and 
supposed to be ordinary bacteria of a genus Fusiformis or Fusobacterium may be its 

Leptospira Noguchi. The cells tightly coiled. L. icterohaemorrhagiae is the agent 
of infectious jaundice. L. icteroides, isolated by Noguchi in South America, sup- 
posedly from cases of yellow fever, is perhaps the same thing: it is now known that 
yellow fever is caused by a virus. It was in pursuing in Africa his study of yellow 
fever that Noguchi lost his life by this disease (Flexner, 1929; Eckstein, 1931). 


Myxophykea Wallroth 1853. 

Myxophyceae Stizenberger 1860. 

Division (of Class Algen) Pkycochromaceae and order Gloiophyceae Rabenhorst 

Krytog.-Fl. Sachsen 1: 56' (1863). 
Cyanophyceae Sachs Lehrb. Bot. ed. 4: 248 (1874). 

OrAtx Cyanophyceae or Pkycochromaceae yicNdLhm]o\iT. oi'Qot. 15: 340 (1877). 
Schizophyceae Cohn 1879, not suborder Schizophyceae Rabenhorst Deutschland's 

Kryptog.-Fl. 2, Abt. 2: 16 (1847). 

30 ] The Classificatio7i of Lower Organisms 

Order Schizophyceae Schenck in Strasburger et al. Lehrb. Bot. 1894. 

Class Schizophyceae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 
la: iii (1900). 

Class Archiplastideae Bessey in Univ. Nebraska Studies 7: 279 (1907). 

Class Cyanophyceae Schaffner in Ohio Naturalist 9: 446 (1909). 

Class Myxophyceae G. M. Smith (1918). 

Subclass Myxophyceae Setchell and Gardner in Univ. California Publ. Bot. 8, 
part 1: 3 (1919). 

Cyanophyta Steinke ( 193 1 ) . 

Stamm Cyanophyta Pascher in Beih. Bot. Centralbl. 48, Abt. 2: 330 (1931). 

Mychota most of which live by phytosynthesis of primitive or typical character, 
many of them, and most of the saprophytic and chemosynthetic organisms included 
with them, being of the form of sheathed filaments. 

This is primarily the group of the blue-green algae. Blue-green algae are familiar 
things as forming dark scums in water and on wet surfaces. Rabenhorst (1863) ap- 
pears first to have recognized them as a group definitely distinct from green algae; 
he named most of the recognized families. Revisions by Thuret (1875), Bomet and 
Flahault (1886-1888), and Gomont (1892) failed to provide a satisfactory system 
of the group; Kirchner's revision (in Engler and Prantl, 1898) is the accepted system. 

One of the important contributions of Cohn was his suggestion that the bacteria 
and blue-green algae belong together. He emphasized this view by mingling the 
genera of the two groups in two new groups, "tribes," named in effect slime-formers 
and thread-formers (1875). In this he went too far; but some of the arrangements 
which he suggested appear natural. Beggiatoa, the type of order Thiobacteria of 
Migula, appears to be a variant of the common blue-green alga Oscillatoria, differing 
from it in living by chemosynthesis. Most of the so-called iron bacteria, family 
Chlamydobacteriaceae of Migula, fall readily into scattered places among the blue- 
green algae. Only the genus Sphaerotilus remains at loose ends. It is credibly reported 
to produce cells swimming by means of flagella; no proper blue-green algae do this. 

A variety of purple bacteria — bacteria, that is, which contain a red pigment — 
have been discovered from time to time. Engelmarm (1888) observed that they swim 
toward the light, and convinced himself that they live by photosynthesis. Van Niel 
confirmed this, and showed that photosynthesis is in this group of a peculiar character; 
it requires the presence of reducing agents and does not release oxygen. This type of 
photosynthesis appears, in fact, to represent a stage of the evolution of typical photo- 
synthesis; the group in which it occurs appears to represent the ancestry of the 
typical blue-green algae. The poorly known green bacteria appear to belong with 
the purple bacteria. 

Various members of this class have been proved capable of fixing nitrogen (Sisler 
and ZoBell, 1951; Williams and Burris, 1952). 

Four orders may be distinguished: 

1. Possessing a red ("purple") intracellular 
pigment, or a green pigment not masked by 
others Order 1 . Rhodobacteria. 

l.With green pigment masked by others, or 

2. Producing cells with flagella; non-pig- 
mented sheathed filaments not accumu- 
lating ferrugineous matter Order 2. Sphaerottlalea. 

Kingdom Mychota [ 31 

2. Never producing cells with flagella. 

3. Cells dividing in more planes than 

one, growing to full size before re- 
dividing; unicellular or colonial, not 

filamentous Order 3. Coccogonea. 

3. Cells dividing in one plane, and 

accordingly producing filaments; 

exceptional examples reproducing 

by budding (unequal division) or 

by repeated division into minute 

spores Order 4. Gloiophycea. 

Order 1. Rhodobaeteria Molisch Purourbakterien 27 (1907). 

Rods, spheres, and spirals, solitary or colonial, with red or green pigment, per- 
forming in the presence of light and reducing substances a sort of photosynthesis in 
which no oxygen is released. 

These organisms have generally been included in Thiobacteria, but do not include 
Beggiatoa, the type of that order. Molisch divided them into two families, Thiorho- 
daceae, aerobic, accumulating granules of sulfur, and Athiorhodaceae, microaero- 
philic or anaerobic, not accumulating granules of sulfur. The green bacteria are to be 
placed as a third family. The names originally applied to the families are not tenable. 

Family 1. Chromatiacea (Migula) nomen familiare novum. Subfamily Chro- 
MATiACEAE Migula. Family Rhodobacteriaceae Migula; Family Thiorhodaceae 
Molisch; the family does not include genera with corresponding names. Purple bac- 
teria, aerobic, accumulating granules of sulfur. Chromatium Perty includes the or- 
ganism of foul waters which was originally named Monas Okenii. It is a plump rod, 
often bent, sometimes exceeding \0\Ji in length, monotrichous or lophotrichous. There 
are a dozen other genera, rods, spheres, and spirals [Thio spirillum., which belongs 
here, is to be distinguished alike from Spirillum, Thiospira, and Rho do spirillum), 
solitary or colonial, motile or non-motile. Most of them were discovered by Wino- 

Family 2. Rhodobacillacea nom. nov. Family Athiorhodaceae Molisch. Molisch 
named in this family a genus Rhodobacterium, but the name Rhodobacteriaceae had 
already been applied by Migula to the preceding family. Purple bacteria, anaerobic, 
not accumulating granules of sulfur. Molisch discovered all known members of the 
present family. The method of culture was to place a mass of organic matter, for 
example an egg, in the bottom of a cylinder of water (the original account specified 
water of the River Moldau), cover the surface with oil, place in a north window, and 
wait several weeks. This method yielded organisms which were assigned to seven 
genera. Those of spiral form are Rhodospirillum. All others are by van Niel treated 
as a single genus, which may be called Rhodobacillus Molisch {Rhodopseudomonas 
van Niel). 

Family 3. Chlorobiacea nom. nov. Family Chlorobacteriaceae Geitler and Pascher 
ex van Niel in Bergey's Manual ed. 6: 869 (1948). Geitler and Pascher (in Pascher 
Siisswasserfl. Deutschland, 1925) did not place this group in a definite category 
and name it unequivocally: they called it Cyanochloridinae or Chlorobacteriaceae. 
Minute spherical or elongate cells with a green pigment different from typical 
chlorophyll, anaerobic, non-motile, producing irregular or regular gelatinous colonies. 
Chlorobium, Pelodictyon, Clathro Moris, with a half a dozen known species. Certain 


The Classification of Lower Organisms 

Fig. 4. — Coccogonea: a, Chroococcus sp.; b, C, Achromatiuni oxalijerum. Gloio- 
phycea: d, Oscillatoria splendida; e, Phormidium sp.; f, Beggiatoa sp.; g, Chamae- 
siphon incrustans; h, Anabaena inacqualis; \, Cylidrospcrmum majus; j, Chlarnydo- 
thrix ochracea; k, 1, m, Clonothrix fusca after Kolk (1938); n, Dermocarpa protea 
after Setchell and Gardner ( 1919); o, Crenothrix polyspora after Kolk ( 1938). All 
X 1,000. 

Kingdom Mychota [33 

organisms of this group, occurring in symbiotic combinations with larger bacteria or 
with protozoa, have been named as additional genera; one of these is Chlorobacterium 
Lauterborn, but the name is a later homonym. 

Order 2. Sphaerotilalea nom. nov. 

Order Desmobactcrialcs Pribram in Jour. Bact. 18: 376 (1929); there is no cor- 
responding generic name. 

Cells colorless, elongate, in sheathed filaments which branch freely in the manner 
called "false": the cells divide strictly in one plane; those at a distance from the tip 
may so multiply as to break the continuity of the series by pushing a growing point 
laterally out of the sheath. The cells may escape from the filaments, become lophotri- 
chous, and function as swarm spores. There is a single family: 

Family Sphaerotilacea [Sphaerotilaceae] Pribram 1. c. There is probably only 
one species, Sphaerotilus natans Kiitzing {Cladothrix dichotoma Cohn). It is found 
as minute gelatinous colonies fioating on stagnant water; cells 2-4^ in diameter. 

Order 3. Coccogonea [Coccogoneae] (Thuret) Campbell Univ. Textb. Bot. 84 
Tribe Chroococcaceac [Coccogoneae) Thuret in Ann. Sci. Nat. Bot. ser. 6, 

1: 377 (1875). 
Subclass Coccogoneae Engler in Engler and Prantl Nat. Pfianzenfam. I Teil, 

Abt. la: iii (1900). 
Order Coccogonales Atkinson 1903. 

Orders Chroococcales and Entophysalidales Geitler in Pascher et al. Siisswasserfl. 
Deutschland 12: 52, 120 (1925). 
Cells solitary or colonial, not filamentous, never flagellate; mostly of blue-green 
color and living by photosynthesis. 

Kirchner (in Engler and Prantl, 1898) placed here two families, Chroococcaceac 
and Chamaesiphonaceae, but the second belongs to the following order. A proper 
second family includes the colorless organisms of genus Achromatium. 

Family 1. Chroococcacea [Chroococcaceac] (Nageli) Rabenhorst Kryptog.-Fl. 
Sachsen 1:69 (1863). Order Chroococcaceac Nageli Gatt. einzell. Alg. 44 (1849). 
Unicellular or colonial blue-green algae. Chroococcus, Gloeocapsa, Merismopedia, 
Coelosphaerium, Gomphosphaeria, etc., occur as plankton or as masses on damp 
surfaces or the bottoms of bodies of water. Certain species occur as symbionts or 
parasites within the cells of the green algae Glaucocystis and Gloeochaete. The re- 
sulting bodies, having the color of blue-green algae with the structure of green 
algae, resisted classification until Geitler (1923) explained their nature. 

Family 2. Achromatiacea [Achromatiaceae] Buchanan. Cells solitary, large, ellip- 
soidal, without flagella, non-pigmented; protoplasm alveolar, with or without 
granules of sulfur in the alveoli. Half a dozen species have been described; Bersa 
(1920) was probably correct in reducing all to the original one, Achromatium 
oxaliferum Schewiakoff. It occurs on mud under still waters rich in organic matter. 

Order 4. Gloiophycea [Gloiophyceae] Rabenhorst Kryptog.-Fl. Sachsen 1 : 56 

Tribe Nostochineae {Hormonogoneae) Thuret in Ann. Sci. Nat. Bot. ser. 6, 
1: 377 (1875). 

34 ] The Classification of Lower Organisms 

Family Hormogoneae Bomet and Flahault in Ann. Sci. Nat. Bot. ser. 7, 3 : 337 

Subclass Hormogoneae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, 

Abt. la: iii(1900). 
Order Hormogoneae Campbell Univ. Textb. Bot. 84 (1902). 
Order Hormogonales Atkinson 1905. 
Blue-green algae whose cells divide predominantly in one plane, so that filaments 
are produced, together with related colorless organisms. 

So far as cell division is strictly in one plane, any branching of the filaments is of 
the type called "false": it occurs by breaks in the continuity of the series of cells, 
followed by the outgrowth, beside the original series, of the newly formed tips. In 
some members of the group, however, the cells are not strictly confined to division in 
one plane, with the result that "true" branching is possible. There are a few appar- 
ently derived examples in which cell division takes place freely in all planes. 

Many of these algae produce spores of the type called arthrospores by the direct 
conversion of normal cells into thick-walled resting spores. Many (almost but not 
quite exactly the same ones which produce arthrospores) produce peculiarly differ- 
entiated cells called heterocysts (the word means "different cells"). These are en- 
larged thick-walled cells with colorless contents; their most obvious function is to 
furnish breaking points for the filaments. They are believed to be variants of the 
arthrospores; they have been seen to germinate and give rise to normal filaments. 
Ten families may be distinguished as follows: 
1. Cells dividing strictly in one plane; branch- 
ing none or of the "false" type. 

2. The filaments not branching nor taper- 
ing nor producing spores or heterocysts. 
3. Filaments elongate. 

4. Pigmented, blue-green Family 1. Oscillatoriacea. 

4. Colorless organisms accumu- 
lating sulfur Family 2. Beggiatoacea. 

3. Filaments reduced to single cells 

which reproduce by budding Family 3. Chamaesiphonacea. 

2. Filaments branching or tapering, or pro- 
ducing spores or heterocysts, or showing 
several of these characters. 
3. Filaments not tapering. 

4. Filaments not branching Family 4. Nostocacea. 

4. Filaments branching. 

5. Blue-green algae mostly 

producing heterocysts Family 5. Scytonematacea. 

5. Minute colorless filaments 

without heterocysts Family 6. Chlamydotrichacea. 

3. Filaments tapering Family 7. Rivulariacea. 

1. Cells dividing in more planes than one, usual- 
ly after a preliminary filamentous phase. 
2. Pigmented, blue-green. 

3. Producing extensive filaments with 

heterocysts Family 8. Sirosiphonacea. 

Kingdom Mychota [35 

3. Filaments more or less reduced, re- 
producing by minute spores (gon- 
idia) formed by repeated division 

in all planes Family 9. Pleurocapsacea. 

2. Colorless; filamentous, reproducing by 

gonidia Family 10. Crenotrichacea. 

Family 1. Oscillatoriacea [Oscillatoriaceae] Harvey 1858. Blue-green algae con- 
sisting of unbranched filaments, not tapering, without spores or heterocysts; mostly 
actively motile by mechanisms as yet unknown. In the commonest genus, Oscillatoria, 
the filaments are straight and lack sheaths. Lyngbya and Phormidium produce 
sheathed filaments, in the latter genus very slender. Microculeus and Hydrocoleum 
have more than one filament in each sheath. In Arthrospira and Spirulina the fila- 
ments are coiled; those of Spirulina are not visibly septate, and are said to be uni- 

Family 2. Beggiatoacea [Beggiatoaceae] Migula 1895. Beggiatoa Trevisan includes 
slender colorless filaments, actively writhing, containing granules of sulfur, found 
in foul waters and sulfur springs. The species were originally included in Oscillatoria. 
Winogradsky (1887) showed that they live by chemosyn thesis, and discovered the 
related genera Thiothrix and Thioploca. From the time of these discoveries, these 
organisms were construed as bacteria of an order Thiobacteria. Under the current 
hypri thesis that chemosynthesis is a derived character, we are free to believe that the 
position originally assigned to the species of Beggiatoa was the natural one. 

Family 3. Chamaesiphonacea [Chamaesiphonaceae] Borzi 1882. Order Chamaesi- 
pkonales Smith Freshw. Alg. 74 (1933). The only genus is Chamaesiphon, minute 
organisms epiphytic on freshwater plants. The ellipsoid cells are attached at one 
end and are enclosed in tenuous sheaths. They reproduce by transverse division, which 
cuts loose small cells from the free ends. By the time two or three such cells are 
produced, the sheath is ruptured at the free end, and the small cells drift away to 
repi educe the organism elsewhere. 

Family 4. Nostocacea [Nostocaceae] (Nageli) Rabenhorst Kryptog.-Fl. Sachsen 
1: 95 (1863). Order Nostocaceae Nageli 1847. Of this family the most familiar genus 
is Nostoc, seen as gelatinous bodies, usually globular, green, blue-green, yellow, or 
brown, of sizes from barely visible to the naked eye up to 10 cm. or more in diameter, 
in fresh water or on damp earth. Under the microscope, these bodies or colonies are 
seen to consist of myriad crooked and tangled filaments of bead-like cells imbedded 
in a gelatinous matrix. Heterocysts are always, and spores usually, present. 

If in water one finds filaments of much the same structure as those of Nostoc, 
but comparatively short, straight, and free or at least not in definite colonies, these 
represent the genus Anabaena. Filaments floating on water, with cylindrical spores 
not confined to the ends of the filaments, are Aphanizomenon. Filaments each with 
one heterocyst and one spore at one end are Cylindrospermum. 

Family 5. Scytonematacea [Scytonemataceae] Rabenhorst op. cit. 106. Members 
of this family produce heavily sheathed filaments like those of Lyngbya, with the 
difference that heterocysts are usually present. The multiplication of the cells of a 
filament may produce the result that the cell next to a heterocyst is driven out of line 
and forced obliquely through the sheath. With further growth, the file of cells ending 
in one which was forced out of line may appear to be the main axis of a system of 
branches, while the original summit of the filament appears to be a lateral branch. 
The description of "false" branching thus given applies particularly to Tolypothrix. 

36 ] The Classification of Lower Organisms 

In Scytonema, the pressure of multiplying cells causes waves of the filament to break 
laterally through the sheath and produce branches in pairs. Plectonema branches 
like Tolypothrix but has no heterocysts. 

Family 6. Chlamydotrichacea [Chlamydotrichaceae] Pribram in Jour. Bact. 18: 
377 (1929). Aquatic organisms consisting of colorless cylindrical cells in sheathed 
filaments, without heterocysts but exhibiting false branching, the sheaths of young 
filaments thin and colorless, those of older ones thick and yellow to brown. Chlamydo- 
thrix ochracea Migula was intended as a new name for Leptothrix ochracea Kiitzing, 
but the entity to which it is believed to apply is totally different from the one to which 
the latter name was applied above. Chlamydothrix is a filament of definite cells 
about 1 ^ in diameter. The only other definitely characterized species of this family is 
Clonothrix fusca Roze, the cells about 2^ in diameter, those near the tips of the fila- 
ments dividing repeatedly (always in one plane) to produce spherical non-motile 
gonidia (Kolk, 1.938). 

Family 7. Fdvulariacea [Rivulariaceae] Rabenhorst op. cit. 101. The filaments 
include heterocysts and exhibit the false branching of Tolypothrix; the outgrowth 
of the filament below each heterocyst gives to the original terminal part the appear- 
ance of a branch of which the heterocyst is the basal cell. The ends of the filaments 
become attenuate and colorless. In Calothrix the filaments are mostly solitary; in 
other genera they remain together in gelatinous colonies. Rivularia is without spores; 
in Glocotrichia there is a large cylindrical spore next to each heterocyst. 

Family 8. Sirosiphonacea [Sirosiphonaceae] Rabenhorst op. cit. 114. Family 
Stigonemataceae Kirchner 1898. This family takes its name from the ancient generic 
name Sirosiphon Kiitzing 1843, which turned out to be identical with Stigonema 
Agardh 1824. The cells divide at first in one plane and produce filaments. Presently 
they exhibit a capacity to divide in other planes, and may produce true branches or 
multiseriate filaments or both. Heterocysts and spc^es are generally produced. 

Family 9. Pleurocapsacea [Pleurocapsaceae] Geitler in Pascher et al. Siisswasser- 
Fl. Deutschland 12: 124 (1925). This group was formerly included in Chamaesi- 
phonacea, but it appears probable that Chamae siphon is related to Oscillatoria, and 
the present group to Stigonema. Most of the Pleurocapsacea are marine, epiphytic 
on seaweeds. Their apparently typical behavior, as exemplified by Hyella and 
Radaisia, consists of the production of branching filaments whose terminal eel's be- 
come enlarged, after which their contents undergo division in many planes to produce 
numerous minute spores called gonidia. In Pleurocapsa and Xenococcus there is no 
filamentous phase; the gonidium gives rise to a cluster of cells all of which produce 
gonidia. In Dermocarpa the gonidium gives rise to a single vegetative cell which 
divides only to produce gonidia. 

Family 10. Crenotrichacea [Crenotrichaceae] Hansgirg. This family includes the 
single known species Crenothrix polyspora Cohn, one of the traditional iron bacteria. 
There is every appearance that it is a colorless variant of the Pleurocapsacea. A germi- 
nating gonidium gives rise to an unbranched filament of cells, about 2^ in diameter, 
in a sheath which is at first thin and colorless, later becoming thicker and discolored 
by ferric oxide. Some cells may burst from the free end of the sheath as macrogonidia. 
Others may begin to divide lengthwise. These may at first grow before re-dividing, 
and may swell the sheath to a fusiform or trumpet-like shape. By further division 
they produce numerous microgonidia, which may sift out of the sheath or be re- 
leased by its decay. 

Such are the Mychota, the organisms which may properly be characterized as 
lacking nuclei. 

Chapter IV 

Kingdom !l. PROTOCTISTA Hogg 

Regne Psycho diaire, Psychodies, Bory de Saint Vincent Diet. Class Hist. Nat. 8: 
246 (1825), 14: 329 (1828). 

Kingdom Protozoa Owen Palaeontology 5 (1860), not class Protozoa Goldfuss 

Regnum Primigenium seu Protoctista Hogg in Edinburgh New Philos. Jour. n.s. 
12: 223 (1860). 

Kingdom Acrita or Protozoa Owen Palaeontology ed 2: 6 (1861). 

Kingdom Primalia Wilson and Cassin in Proc. Acad. Nat. Sci. Philadelphia 1863: 
117 (1864). 

Kingdom Protista Haeckel Gen. Morph. 2: xix (1866). 

Kingdom Protobionta Rothmaler in Biol. Centralbl. 67: 243 (1948). 

Nucleate organisms other than Plantae and Animalia: the marine algae and the 
fungi and protozoa. Amiba diffluens may be construed as the standard. 

The name Protista, of Haeckel, is the most familiar among those which have been 
applied to the kingdom here to be discussed, but it is not the earliest. Among fol- 
lowers of Cuvier, the animal kingdom consisted necessarily of four branches. Presum- 
ably, it was this tradition that induced Owen to refer the Infusoria and Amorphozoa 
(sponges) to a separate kingdom, which he called Protozoa. A year later, Owen pub- 
lished an alternative name for this kingdom; but Hogg had already published modi- 
fications of two of Owen's names, Protoctista and Amorphoctista(KTi^co,to establish, 
create), for the reason that names in -zoa appeared inappropiiate to groups excluded 
from the animal kingdom. 

The limits here given to the kingdom Protoctista were proposed by the present 
author (1938, 1947). They have been accepted, with exception in a single significant 
point, by Barkley (1939, 1949) and Rothmaler (1948). 

It is assumed that the evolutionary origin of the Protoctista consisted of the evolu- 
tionary origin of the nucleus, and that all nuclei are essentially the same thing. Kofoid 
(1923) insisted that enduringly viable nuclei originate among protozoa, as among 
plants and animals, regularly by mitosis, never by binary or multipe fragmentation, 
nor by aggregation of stainable granules. He did not recognize the nucleus as essen- 
tially a device for sexual reproduction. Several considerable groups of protozoa, how- 
ever, which Kofoid listed as not known to reproduce sexually, have been found to 
do so. Here, then, it is maintained that all nuclei, in this kingdom as among plants 
and animals, are the same thing; and that the nucleus is essentially a device for sexual 
reproduction, that is, for processes of reproduction which involve always one act of 
meiosis and one of karyogamy, and which produce Mendelian heredity as an effect. 

Photosynthesis is believed to have evolved only cnce. As it occurs both among non- 
nucleate and nucleate organisms, the nucleus is believed to have evolved in organisms 
living by this function. The closest approach between non-nucleate and nucleate or- 
ganisms is believed to be between the blue-green algae and the primitive red algae 
(Smith, 1933; Tilden, 1933). Thus it appears that the original nucleate organisms 
were not capable of swimming by means of flagella. Flagella appear to have evolved 
in unicellular nucleate photosynthetic organisms as a device for dissemination (Bes- 

38 ] The Classification of Lower Organisms 

sey, 1905). The flagella of nucleate organisms are not homologous with those of 
bacteria; they are much larger and of much more complicated structure. 

The origin of flagella was apparently associated with a simplification of the system 
of photosynthetic pigments, by the loss of chromoproteins, leaving systems of chloro- 
phylls and carotinoids. The association of these two courses of evolution may have 
been merely coincidental; Tilden suggested the idea that the loss of chromoproteins 
may have been occasioned by increasing illumination of the waters of the face of the 

Organisms of the body type of solitary walled cells, having chlorophylls and caro- 
tinoid pigments but not chromoproteins, and producing flagellate reproductive cells, 
appear to have undergone radiating evolution, producing a wide variety of types of 
organisms, distinguished by different specific chlorophylls and carotinoids, different 
types of flagella, and different specific metabolic products. The types of flagella oc- 
curring in nucleate organisms are here particularly to be noted. 

Loeffler (1889), in the original publication of the standard method of staining the 
flagella of bacteria, remarked that he had applied this method also to certain larger 
organisms. He found that the flagellum of Manas bears numerous lateral appendages, 
and that the cilia of a certain infusorian bear solitary terminal appendages. Loeffler's 
method is difficult, and has not been much used. Fischer (1894) used it and coined 
terms, Flimmergeisseln and Peitschengeisseln, designating structures of the respective 
types seen by Loeffler. Petersen (1929), having applied Loeffler's method to a reason- 
able variety of flagellates, introduced refinements of terminology. Flagella of the 
type of the larger flagellum of Monas (the organism bears also a minute simple flagel- 
lum) became allseitswendige Flimynergeisseln; those of Euglena, which bear a single 
file of appengages, became einseitswendige Flimmergeisseln. 

Deflandre ( 1934) devised a different method for seeing the appendages on flagella, 
and substituted, for the Teutonisms just quoted, French terms based on Greek. These 
may be Anglicised as follows. ( 1 ) The acroneme flagellum bears a single terminal 
appendage. The flagellum without appendages is said to be simple; so far as it ap- 
pears among nucleate organisms, it appears to be a variant of the acroneme type. 
(2) The pantoneme flagellum bears appendages on all sides. (3) The pantacroneme 
flagellum bears both terminal and lateral appendages. It is a rarity, known only in 
the collared monads, and may be supposed to be a variant of the pantoneme type. 
(4) The stichoneme flagellum bears a single file of appendages. 

The point in which Barkley and Rothmaler take exception to the limits here given 
to kingdom Protoctista is this, that they include in this kingdom the green algae. In 
the present work, scant attention is given to organisms whose plastids are bright 
green, containing chlorophylls a and b, carotin, and xanthophyll, and no other pig- 
ments; whose motile stages have acroneme flagella, more than one (usually two), 
and equal; and which produce essentially pure cellulose, true starch, and sucrose. 
These organisms represent the undoubted evolutionary origin of the higher plants; 
a classification which attempts to represent nature includes them necessarily in the 
plant kingdom. 

Rothmaler set up a system of only four phyla, being the red organisms, basically 
without flagella; those which are typically yellow to brown, having pantoneme flagel- 
la; those with acroneme flagella, including the green algae; and the euglcnid group, 
which have stichoneme flagella. The non-pigmented Protoctista were distributed 
among these groups. The system appears unsound by the fact that large blocks of 
non-pigmented organisms are placed where only portions of them belong. 

Kingdom Protoctista [ 39 

In the present work, a less symmetrical system of phyla is offered. Its basis is an 
ingenuous system of red algae, brown algae, fungi, and the four traditional groups of 
protozoa; this has been radically modified in view of the great accumulation of 
knowledge subsequent to the formulation of these groups. The phylum Pyrrhophyta 
as here limited is tentative; the phylum Protoplasta, marked only by negative char- 
acters, amounts to a dumping ground for groups whose relationships are altogether 

1. Living by photosynthesis, which takes place 
in plastids containing red or blue chromo- 
protein pigments; never producing flagellate 

cells Phylum 1. Rhodophyta. 

1. Without chromoprotein pigments. 

2. Typically living by photosynthesis, 
brown, yellow, or green in color. 

3. Producing flagellate cells each with 
one pantoneme or pantacroneme 
flagellum, often with additional 

acroneme flagella Phylum 2. Phaeophyta. 

3. Producing flagellate cells whose fla- 
gella are never pantoneme or pan- 
tacroneme, often stichoneme Phylum 3. Pyrrhophyta. 

2. Dependent; motile cells with acroneme 

flagella or cilia, or amoeboid, or none. 

3. Not producing cilia, i. e., structures 

of the nature of acroneme flagella, 

numerous and widely distributed 

on the surfaces of the cells. 

4. Cells walled in the vegetative 

5. Producing motile cells 
with single posterior fla- 
gella; bodies mostly with 

tapering rhizoids Phylum 4. Opisthokonta. 

5. Producing no motile cells; 

bodies filamentous Phylum 5. Inophyta. 

4. Cells not walled in the vegeta- 
tive condition. 
5. Mostly predatory, flagel- 
late or amoeboid or with 

flagellate or amoeboid stages Phylum 6. Protoplasta. 

5. Parasitic in animals, pro- 
ducing flagellate cells only 

as rare exceptions Phylum 7. Fungilli. 

3. With cilia .Phylum 8. Ciuophora. 

Chapter V 

Phylum 1. RHODOPHYTA Wettstein 

Order Floridees Lamouroux in Ann. Mus. Hist. Nat. Paris 20: 115 ( 1813) . 

Florideae C. Agardh Synops. Alg. Scand. xiii (1817). 

Order pLORroEAE C. Agardh Syst. Alg. xxxiii (1824). 

Division (of order Algae) Rhodospermeae Harvey in Mackay Fl. Hibern. 160 

Class Heterocarpeae Kiitzing Phyc. Gen. 369 (1843). 

Class Florideae J. Agardh Sp. Alg. 1: v (1848). 

Rhodophyceae Ruprecht in Middendorff Sibirische Reise 1, Part 2: 200 (1851). 

Stamm Florideae Haeckel Gen. Morph. 2: xxxiv (1866). 

Phylum RHODOPHYTA Wettstein Handb. syst. Bot. 1: 182 (1901). 

Division Rhodophyceae Engler Syllab. ed 3: 18 fl903). 

Phylum Carpophyceae Bessey in Univ. Nebraska Studies 7: 291 (1907). 

Phylum Rhodophycophyta Papenfuss in Bull. Torrey Bot. Club 73: 218 (1946). 

Definitely nucleate organisms {Porphyridium and Prasiola doubtfully so); with 
few exceptions living by photosynthetic processes involving red and blue pigments 
(phycocyanin and phycoerythrin) as well as green and yellow (chlorophylls a and d 
and carotinoids); not producing true starch, and producing cellulose only in small 
quantity, the cells walled chiefly with modified carbohydrates which tend to become 
gelatinous; never producing flagellum-bearing cells, but sometimes producing cells 
which move in water without the use of definite organelles. 

Tilden (1933) and Smith (1933) are authority for placing the red algae next to 
the blue-green algae, thus suggesting the inference that they include the most primi- 
tive of nucleate organisms. The resemblances between blue-green and red algae 
are in the following points. Both groups possess, along with the chlorophylls and 
carotinoids usual in photosynthetic organisms, other pigments, both blue and red. To 
these pigments as found in both groups, the same names, phycocyanin and phycoery- 
thrin, are applied; they are not, however, the same chemical species (Kylin, 1930). 
Neither group produces true starch; carbohydrate is stored as substances of the 
general nature of dextrin or glycogen (occuring in the red algae as solid granules 
called floridcan starch). Both groups produce cellulose only in scant quantities 
(Miwa, 1940; Kylin, 1943); the cell walls consist chiefly of materials, of the general 
nature of carbohydrates, which tend to become gelatinous. They share the negative 
character of never producing flagella, and the positive one of producing cells which 
call move actively upon surfaces, without motor organelles, by a mechanism as yet 
unknown (Roscnvinge, 1927). 

The phylum is divisible into two classes: 

1. Cells of most examples each with one central 
plastid, without protoplasmic interconnec- 
tions, in aggregates of indefinite extent or or- 
ganized as filaments or thalli with intercalary 
growth; zygotes producing spores directly by 
division Class 1. Bangialea. 

Phylum Rhodophyta [41 

1. Cells with protoplasmic interconnections, 
containing except in the lowest examples sev- 
eral parietal plastids, organized as filaments 
with apical growth, the filaments usually 
massed as thalloid bodies; zygotes giving rise 
to spores indirectly Class 2. Heterocarpea. 

Class 1. BANGIALEA (Engler) Wettstein 

Subclass Bangioideae de Toni Sylloge Algarum 4: 4 (1897). 

Subclass Bangiales Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 2 : 

ix (1897). 
Class Bangiales Wettstein Handb. syst. Bot. 1: 187 (1901). 
Class Bangioideae and orders Bangiales and Rhodochaetales Bessey in Univ. Ne- 
braska Studies 7: 291 (1907). 
Class Bangieae SchaflFner in Ohio Naturalist 9: 448 (1909). 
Protoflorideae Rosenvinge in Mem. Acad. Roy. Sci. Lett. Danemark, ser. 7, 

Sciences 7: 55 (1909). 
Abtheilung (of Stamm Rhodophyta) Bangiineae Pascher in Beih. bot. Centralbl. 

48, Abt. 2: 328 (1931). 
Subclass Protoflorideae Smith Freshw. Algae 120 (1933). 

Red algae (exceptionally green or of other colors), the cells with solitary central 
plastids (exceptionally with multiple parietal plastids), lacking protoplasmic inter- 
connections, in irregular colonial masses or forming filaments or thalli with intercalary 
growth; the zygote produced in sexual reproduction dividing to produce spores 

The group is of one order, five families, about fifteen genera; the number of 
known species is about eighty. 

Order Bangiacea [Bangiaceae] Nageli 1847. 
Characters of the class. 

1. Cells forming irregular aggregates Family 1. Porphyridiacea. 

1. Cells forming filaments or thalli. 

2. Vegetative cells becoming spores with- 
out dividing Family 2. Rhodochaetacea. 

2. Vegetative cells undergoing division to 
produce spores. 

3. Organisms red, purplish, etc Family 3. Porphyrea. 

3. Organisms green Family 4. Schizogoniacea. 

2. Spores formed solitary in special cells Family 5. Compsopogonacea. 

Family 1. Porphyridiacea [Porphyridiaceae] Kylin in Kungl. Fysiog. Sallsk. 
Forhandl. 7, no. 10: 4 (1937). Order Porphyridiales Kylin 1. c. The only well known 
species is Porphyridium cruentum (C. Agardh) Nageli (1849). It is widely dis- 
tributed in damp climates, forming extensive red patches like blood on damp earth or 
stone. The spherical cells are reported as varying widely in diameter (5-24^), and 
Geitler (1932) and Kylin (1937) have distinguished additional species. 

Porphyridium has been classified among blue-green, red, and green algae. Lewis 
and Zirkle (1920) found in each cell a central red plastid, occupying most of its 
volume, and having rays extending to the cell membrane. Within the plastid there is 


The Classification of Lower Organisms 

Fig. 5. — a, Porphyra laciniata, thallus x 1/2. b-g, Porphyra tenera after Ishikawa 
(1921); b, cells; c, cell dividing to produce sperms; d, sperms; e, fertilization; 
f, "carpospores," i.e., cells produced by division of the zygote; g, stages of nuclear 
division x 2,000. h, i, Porphyra umbilicaris after Dangeard ( 1927); h, fertilization; 
i, stages of nuclear division x 2,000. All figures x 1,000 except as noted. 

Phylum Rhodophyta L 43 

a moderately large stainable granule; outside the plastid, a single additional granule 
can usually be found. When a cell is to divide, the granules break up into consid- 
rable numljers of smaller ones, some of which become organized as a system of strands 
forming an irregular network on the surface of the plastid. The protoplast, the 
network, and the plastid undergo constriction; the processes by which the daughter 
cells return to the original structure were not clearly seen. Interpretation of these 
observations is difficult. It is possible that the granule outside of the plastid is a 
nucleus of the type of those which have been observed in Bangia and Porphyra. 

Family 2. Rhodochaetacea [Rhodochaetaceae] Schmitz in Engler and Prantl Nat. 
Pflanzenfam. I Teil, Abt. 2: 317 (1896). Family Goniotrichaceae Smith Freshw. 
Algae 121 (1933). Branching filaments, sometimes becoming multiseriate by length- 
wise division, the vegetative cells capable of escaping and functioning as spores. 
Sexual reproduction unknown. Asterocystis, uncommon, in fresh water; the remain- 
ing genera marine, epiphytic on other algae. Goniotrichum. Rhodochaete and Gonio- 
trichopsis, the cells with numerous plastids. 

Family 3. Porphyrea [Porphyreae] Kutzing (1843). Family Porphyraceae Raben- 
horst 1868. Family Bangiaceae (Nageli) Schmitz (in Engler and Prantl, 1896). 
Filaments or thalli of a red or purple color; the cells, in producing spores, may re- 
lease their protoplasts as wholes or may undergo division into many. Rosenvinge 
(1927) observed the active motion of these spores. 

The most important genus is Porphyra; the individuals are thalli up to several 
centimeters in diameter, on rocks or other algae in ocean water along coasts. They 
are called purple lavers, tsu'ai, amanori; they are used as food, for making soup or in 
condiments, and are extensively cultivated in Japan (Tseng, 1944). Bangia is either 
freshwater or marine; in structure it differs from Porphyra in having filementous 
bodies, uniseriate or pluriseriate. 

During nuclear division in Porphyra tenera as described by Ishikawa (1921), polar 
appendages form at both ends of the nucleus, which becomes elongate and appears 
to consist of three strands. The strands break transversely, and each set of three fuses 
into a mass. Dangeard (1927), dealing with Porphyra umbilicaris and Bangia fusco- 
purpurea, observed nuclei 5^ in diameter, each consisting of a karyosome, that is, a 
mass of chromatin, lying in a clear space surrounded by a membrane. In mitosis, the 
membrane and the unstained matter disappear. Polar appendages grow out from the 
karyosome, and their tips become cut off as granules which may be regarded as cen- 
trosomes. The remainder of the karysome becomes organized as two masses, evidently 
chromosomes, connected to the centrosomes by fibers. Each chromosome divides into 
two; the daughter chromosomes move to the centrosomes and fuse with them to form 
karyosomes about which new membranes appear. This description represents a defi- 
nite, if primitive, process of mitosis. 

Sexual reproduction, here where we first encounter it, involves differentiated ga- 
metes. Naked sperms, indistinguishable from spores, move to the surface of other 
cells which function as eggs. A strand of protoplasm grows through the gelatinous 
wall of the egg from the sperm to the egg protoplast, and the protoplast of the sperm 
migrates through the passage thus formed. The zygote divides two or three times, 
producing spores. During the first two divisions, the two masses of chromatin which 
appear are somewhat different in appearance from the vegetative chromosomes 
(Dangeard, op. cit.); it may be supposed that these masses are tetrads and diads, and 
that the divisions are meiotic. Evidently, this is a life cycle of the primitive type, 
in which all cells except the zygotes are haploid. 

44 ] The Classification of Lower Organisms 

Family 4. Schizogoniacea [Schizogoniaceae] Chodat. Family Prasiolaceae West. 
Family Blastosporaceae Wille. Filamemous or thallose algae, freshwater or marine, 
of the structure of Porphyrea, but of a green color; sexual reproduction unknown. 
Kylin (1930) found the pigmentation to be that of green algae rather than of red. 
Copeland (1955) was unable to discern nuclei. The sole genus Prasiola {Schizo- 
gonium represents a stage of development) is of about fifteen species. Setchell and 
Gardner (1920) and Ishikawa (1921) suggested the place in Bangiacea here given to 
this group. 

Family 5. Compsopogonacea [Compsopogonaceae] Schmitz in Engler and Prantl 
Nat. Pflenzenfam. I Teil, Abt. 2: 318 (1896). Family Erythrotrichiaceae Smith 
Freshw. Algae 122 (1933). Filaments, unbranched or branched, uniseriate or pluri- 
seriate, or thalli. Spore-formation is accomplished by the division of a vegetative cell, 
by an oblique wall, into two unequal cells; the protoplast of the smaller is released as 
a spore. Rosenvinge observed the spores of Erythrotrichia cornea to move as far as 
140[i per minute. Sexual reproduction is much as in Porphyrea. Erythrotrichia. 
Erythrocladia. Compsopogon, in fresh water, the cells with numerous parietal plastids. 

Class 2. HETEROCARPEA Kutzing 

Class Heterocarpeae Kiitzing Phyc. Gen. 369 (1843). 

Class Florideae (C. Agardh) J. Agardh Sp. Alg. 1 : v ( 1848). 

Subclass Florideae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 2: 
ix (1897). 

Subclass Euflorideae de Toni Sylloge Algarum 4: 4 (1897). 

Abtheilung (of Stamm Rhodophyta) Floridineae Pascher in Beih. bot. Centralbl. 
48, Abt." 2: 328 (1931). 

As this is the type group of phylum Rhodophyta, most of the synonymy of that 
name applies to this one also. 

Red algae whose bodies consist essentially of filaments growing apically, the cells 
with protoplasmic interconnections, the plastids (except in some of the lowest ex- 
amples) of the form of multiple parietal disks; the filaments commonly compacted 
into cylindrical or thallose bodies; zygotes not dividing to form spores directly, pro- 
ducing spores by budding or indirectly by processes of growth of various degrees of 

In undertaking to describe the varied, and often highly complicated, reproductive 
processes of the typical red algae, one notes that these organisms occur as haploid 
individuals, and that the majority occur as distinct male and female haploid individ- 
uals. Sperms (commonly called spermatia) are minute naked protoplasts released 
from small cells commonly occurring in patches on the surfaces of thalli. The egg 
is called a carpogonium (Schmitz, 1883). It is the terminal cell of a specialized fila- 
ment, the carpogonial filament, and bears a filiform terminal extension, the tri- 
chogyne (Bornet and Thuret, 1867), whose function is to receive the sperms. The 
cell, often diflferentiatcd, from which the carpogonial filament grows, is the support- 
ing cell [Trugzelle). 

In the more primitive members of the class, the zygote gives rise by budding to a 
mass of cells called the cystocarp. The cells of the cystocarp release their protoplasts 
as spores called carpospores. These on germination produce haploid individuals like 
the original ones. The zygote nucleus is the only diploid nucleus in the life cycle; its 
first divisions arc meiotic. 

Phylum Rhodophyta 


In more advanced examples, the first step of development after fertilization con- 
sists of the establishment of protoplasmic contact between the zygote and other cells. 
These may be adjacent cells, reached directly, or distant cells, reached by the out- 
growth of connecting filaments from the zygotes. In the generality of the group, the 
cells with which contact is made give rise to cystocarps producing carpospores; in 
this situation, the cells in question are called auxiliary cells. In some examples, the 
connecting filaments, after making contact with cells called nurse cells, themselves 
give rise to the cystocarps. The carpospores, in all of these more advanced examples, 
give rise to diploid individuals. The diploid individuals are of the same vegetative 

'ymW''^- :^^>^^ 




Fig. 6 — Nuclear phenomena in Polysiphonia violacea after Yamanouchi (1906). 
a, b, c. Stages of mitosis, d, e. Stages of homeotypic division. 

structure as the haploid individuals, but do not produce spermatia, carpogonia, or 
cystocarps. Certain cells, commonly scattered and imbedded in the body, produce 
sets of four spores which are accordingly called tetraspores; these give rise to haploid 

This account means that these algae occur typically in somata of four types: male 
and female haploid individuals; cystocarps, being a preliminary, parasitic, multipli- 
cative phase of the diploid stage (Janet named this stage the carposporophyte; Drew, 
1954); and free-living diploid individuals, reproducing by tetraspores. The produc- 
tion of carpospores and tetraspores by different individuals of identical vegetative 
structure explains the oldest name applied to this class, namely Heterocarpea. 

Understanding of the life cycle of typical Heterocarpea has been reached only by 
much labor and after a certain amount of confusion. The first significant observations 
were by Bornet and Thuret (1867). Schmitz (1883) showed that the zygote makes 
protoplasmic contact with other cells. He supposed that the contact of the zygote 
with an auxiliary cell is a second sexual fusion {Copulation) following upon proper 

46 ] The Classification of Lower Organisms 

fertilization. Oltmanns (1898) disproved this: he showed that the nuclei of auxiliary 
cells are inert, and that the nuclei of carpospores are derived entirely from zygote 
nuclei. Yamanouchi (1906) showed that the chromosome number of carposporic 
individuals of Polysiphonia violacea is 10, and that that of tetrasporic individuals is 
20; and reported much more of the cytology. Centrosomes appear de novo during the 
earlier stages of mitosis, and fade out and disappear during the later stages. The 
mitotic spindle is formed, and the chromosomes take their place upon it, within an 
intact nuclear membrane, which fades out in later stages. In meiosis, which produces 
the nuclei of tetraspores, the tetrads and diads divide within the original nuclear 
membrane, which becomes tetrahedrally lobcd, and then disappears except where 
the haploid groups of chromosomes lie against it, with the result that the membranes 
of the tetraspore nuclei are partly old and partly new. 

There are some 2500 species of Heterocarpea, including comparatively few in 
fresh water, but the majority of the marine algae. Many of them are beautiful; their 
variety and beauty contribute to the pleasure which people find on coasts. Exper- 
ienced naturalists can identify many genera by gross structure, but the systems of 
orders and families based on gross structure, such as those of Kiitzing (1843) and J. 
Agardh (1851-1863), have been found artificial and abandoned. A proper respect 
for the principles of nomenclature makes it necessary, however, to apply many of the 
names used in these systems. Schmitz applied his morphological studies to a classifica- 
tion of the typical red algae as four groups ( 1889) ; Engler ( 1897) made these groups 
definitely orders. Subsequent scholars have found this system sound in principle, but 
have found it necessary, on the basis of studies of additional examples (for example, 
by Kylin, 1923, 1924, 1925, 1928, 1930, 1932; Papenfuss, 1944; Sjostedt, 1926; 
Svedelius, 1942) radically to rearrange the families and genera. At least four orders 
in addition to those of Engler have been proposed but reductions have decreased 
the number currently recognized to six. 

The following key to the orders is a rather considerable modification of those pub- 
lished by Kylin (1932) and Smith (1944). 

l.All free-living individuals haploid; tetra- 
spores not produced, or produced as carpospores. . Order 1. Cryptospermea. 
1. Free-living individuals of two types, the one 
producing gametes (the zygotes giving rise 
to carpospores), the other producing tetraspores. 
2. Without specialized auxiliary cells or 
nurse cells, the lower cells of the carpo- 
gonial filaments, or normal vegetative 

cells, serving as auxiliary cells Order 2. Sphaerococcoidea. 

2. With specialized nurse cells, the carpo- 
spores produced from filaments which 

have made contact with these Order 3. Gelidialea. 

2. With specialized auxiliary cells from 
which the carpogonia develop. 

3. The auxiliary cells being intercalary 
cells in specialized filaments homol- 
ogous with the carpogonial filaments. . . . Order 4. Furcellariea. 
3. The auxiliary cells terminal in fila- 
ments which grow from the support- 
ing cells of the carpogonial fila- 
ments before fertilization Order 5. Coeloblastea. 

Phylum Rhodophyta [ 47 

3. The auxiliary cells originating after 
fertilization as branches of the sup- 
porting cells of the carpogonial 
filaments Order 6. FLORroEA. 

Order 1. Cryptospermea [Cryptospermeae] Kiitzing Phyc. Gen. 321 (1843). 
Order Periblasteae Kutzing op. cit. 387, in part. 
Orders H elmint hoc lade ae J. Agardh Sp. Alg. 2: 410 (1851), Chaetangieae op. 

cit. 456 (1851), and Wrangelieae op. cit. 701 (1863). 
Order Batrachospermaceae'R.ahtnhovstKxy^X.og.-Yl.^dichstn 1: 278 (1863). 
Nemalioninae Schmitz in Flora 72: 438 (1889). 

Order Nemalionales Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 
2: ix (1897). 
Heterocarpea normally without diploid bodies, the carpogonium arising from the 
zygote or from an adjacent cell serving as an auxiliary cell, the carpospores produc- 
ing haploid bodies like the original ones. Certain genera which are exceptional to 
these characters are noted below. Batrachospermum may be regarded as the standard 

In all recent literature, this order is called Nemalionales. Eight families are rec- 
ognized. The forms consisting of mere filaments, Acrochaetium, Rhodochorton, and 
others, are family Acrochaetiacea [Acrochaetiaceae] Fritsch (Family Chantransi- 
aceae Auctt., but Chantransia DC. as originally published included no members of 
this family; Papenfuss, 1945). In the remainder of the order, the filaments are 
differentiated, or, with or without differentiation, organized as bodies of definite 
form, simply cylindrical, branched, or flattened. Fresh-water examples (the only 
fresh-water Heterocarpea) include Batrachospermum, Lemanea, and Thorea. These 
organisms are not red, but bluish, green, or brown. Marine examples include Nemalion 
and Cumagloia. 

In Liagora tetrasporifera and certain other species tetraspores are produced in the 
place of carpospores. Within this genus, then, there has been a change in the time of 
meiosis (which could be established, presumably, by a single mutation) from im- 
mediately after fertilization to the end of the cystocarp stage. 

Galaxaura is a genus of tropical marine algae which are calcified, which is to say 
that they deposit much calcium carbonate in the tissues; they were originally classi- 
fied as corals. They have distinct sexual and tetrasporic stages. Svedelius (1942) as- 
certained their life cycle. Carpospore-bearing filaments arise both from the zygote 
and from other cells, previously undifferentiated, which serve as auxiliary cells. The 
genus has the structure of the present order, and is to be placed here, in spite of ex- 
hibiting in unspecialized form the life cycle of the following orders. 

Order 2. Sphaerococcoidea [Sphaeroccoideae] J. Agardh Sp. Alg. 2: 577 (1852). 
Family Gigartineae Kiitzing (1843). 

Orders Gigartineae and Chaetangieae J. Agardh op. cit. 229, 456 (1851). 
Gigartininae Schmitz in Flora 72: 440 (1889). 
Order Gigartinales Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 

2: X (1897). 
Order Nemastomatales Kylin in Kgl. Fysiog. Sallsk. Handl. n. f. 36, no. 9 : 39 

Order Sphaerococcales Sjostedt in Kgl. Fysiog. Sallsk. Handl. n. f. 37, no. 4: 

75 (1926). 


The Classification of Lower Organisms 

(Legend on bottom of page 49) 

Phylum Rhodophyta [ 49 

This order, in all recent literature called Gigartinales, is a numerous and varied 
one. The bodies are generally erect; they may be cylindrical or flattened, unbranched 
or branched. In some examples, Haliarachnion, Rhodophyllis, Sebdenia, the zygote 
sends out extensive filaments, which make contact with unspecialized cells scattered 
in the body. In other examples, the zygote makes contact with a lower cell of the 
carpogonial filament. In either case, the cells with which contact is made are auxiliary 
cells and give rise to cystocarps; these produce carpospores, and the carpospores pro- 
duce tetrasporic individuals. Certain species of Phyllophora, Gymnogongrus, and 
Ahnfeldtia are exceptional in producing tetraspores in the place of carpospores; these 
species have no free-Uving tetrasporic generation. In these organisms, as contrasted 
with Liagora tetrasporifera, it is believed that this type of Ufe cycle has been estab- 
lished by reduction of a longer one. 

Kylin (1932) assigned twenty families to this order. Gracilaria is a minor source 
of agar agar. Gigartina mammilosa and Chondrus crispus (Irish moss or carageen) 
are well known as yielding a jelly, carageenin, resembling but distinct from agar agar 
(Tseng, 1945). 

Various abnormal growths on red algae have been found to be parasitic red algae, 
almost always on hosts closely related to themselves (Setchell, 1914). To the present 
order belong Gardneriella and its host Agardhiella; Plocamiocolax and its host Plo- 
camium; Gracilariophila and its host Gracilaria (Wilson, 1910). 

Order 3. Gelidialea [Gelidiales] Kylin in Kgl. Svensk. Vetensk.-Akad. Handl. 
63, no. 11: 132 (1923). 
Family Gelidieae Kiitzing (1843). 
Order Gelidieae J. Agardh Sp. Alg. 2: 464 (1851). 
Heterocarpea in which the zygote sends out a single elongate filament which makes 
contact successively with several chains of nurse cells and gives rise to carpospores; 
bodies consisting of branched filaments, the ultimate tips of the lateral branches 
compacted into a firm layer covering a branching body, cylindrical or flattened; the 
surface adjacent to the masses of carpospores pushed out and punctured by pores 
through which the spores escape. 

There is a single family Gelidiea [Gelidieae] Kiitzing ( Family Gelidiaceae Schmitz 
and Hauptfleisch). Such economic importance as the red algae possess lies chiefly in 

Fig. 7 — a, Thallus of Nemalion multifidum x 1. b, c, d^ production of sperms; 
beginning of production of carpospores; and cluster of carpospores of Nemalion 
multifidum after Bornet and Thuret (1867). e, Thallus of Chondrus crispus x 1. 
{, Reproduction of Dudresnaya purpurifera (order Furcellariea or Cryptonemiales) 
after Bornet and Thuret, op. cit. The trichogyne, whose free end with attached 
sperms is seen above, is irregularly twisted below; it leads to the egg (carpogonium); 
connecting filaments, growing from cells below the egg, make contact with auxiliary 
cells at the summits of specialized filaments; each auxiliary cell gives rise to a cluster 
of carpospores. g, Thallus of Delesseria sinuosa x 1. h^ Longitudinal section of 
conceptacle of Polysiphonia nigrescens x 500, after KyUn (1923). The zygote z is 
the fourth and terminal cell of the carpogonial filament whose connection with the 
supporting cell b is not shown; the auxiliary cell a has grown from the supporting 
cell after fertilization. 

50 ] The Classification of Lower Organisms 

this family, and particularly in the genus Gelidium. It is the chief source of agar agar. 
This is the principal material of the cell walls of Gelidium. It is a jelly consisting 
essentially of chains of galactose units, and has the property, that having been melted 
by heat, it does not again become solid until cooled to a much lower temperature. 
Algae containing it have long been used as foods in the orient. Brought into labora- 
tory use by Koch, it has become a necessity in routine bacteriological work. The chief 
source is Japan. 

Kylin construed this order as relatively primitive; but its reproductive processes, 
involving specialized nurse cells, appear less primitive than those of the Sphaerococ- 
coidea. The production of elongate connecting filaments is shared with certain 
examples both of the preceding order and of the following, and the Gelidialea are 
probably derived by specialization from one or the other. 

Order 4. Furcellariea [Furcellarieae] Greville Alg. Brit. 66 (1830). 

Orders Spongocarpeae and Gastrocarpcae Greville op. cit. 68, 157 (1830). 

Order Epiblasteae Kiitzing Phyc. Gen. 382 (1843). 

Orders Cryptonemeae, Dumontieae, Squamarieae, and Corallineae J. Agardh Sp. 

Alg. 2: 'l55, 346, 385 (1851), 506 (1852). 
Cryptoneminae Schmitz in Flora 72: 452 (1889). 

Order Cryptonemiales Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, 
Abt. 2: xi (1897). 

The individuals are crustose or thallose, the thalli cylindrical or flattened, un- 
branched or branched. On the two or three types of individuals of each species, the 
reproductive structures may be scattered or clustered on the surfaces or gathered in 
specialized pits called conceptacles. The eggs are as usual the terminal cells of spe- 
cialized filaments; other filaments, homologous with these but abortive, bear the 
auxiliary cells. After fertilization, the zygote may or may not establish connection 
with a lower cell of the same filaments. Under either circumstance, it sends out fila- 
ments which establish connection with the auxiliary cells, and these send out filaments 
which bear the carpospores. In less specialized examples, the filaments growing from 
the zygote may extend widely through the body; a single one, branching, may reach 
many auxiliary cells. 

Kylin ( 1932) placed nine families here. 

The family Corallinea [Corallineae] Kiitzing (family Corallinaceae Hauck) is one 
of the more specialized. The eggs, and subsequently the carpospores, are clustered in 
conceptacles. In each conceptacle the zygotes, the filaments from them, and the 
auxiliary cells, unite eventually in a single large multinucleate cell from whose mar- 
gins grow the filaments which bear the carpospores. Members of this family have the 
property of accumulating and depositing calcareous material, and were originally 
classified as corals. In modern usage, the term coral means certain lower animals; 
but the coralline algae are associated with them in coral reefs, being indeed, accord- 
ing to Setchell (1926) and other authorities, responsible for the building of the reefs. 
Fossil coralline algae are known from the Ordovician. 

The parasite Callocolax and its host CallophylUs belong to this order; Coreocolax, 
belonging to this order, attacks species of order Floridea. 

The Furcellariea are a numerous group, rather unspecialized, varied almost to the 
extent of a miscellany. They are related to the Sphaerococcoidea, and are believed 
to represent the ancestry of the two following orders, and possibly also of the 

Phylum Rho do phyta [51 

Order 5. Coeloblastea [Coeloblasteae] Kutzing Phyc. Gen. 438 (1843). 
Order Rhodymenieae J. Agardh Sp. Alg. 2: 337 (1851). 
Rhodymeninae Schmitz in Flora 72: 442 (1889). 

Order Rhodymeniales Engler in Engler and Prantl. Nat. Pfllanzenfam. I Teil, 

Abt. 2: X (1897). 

Heterocarpea producing auxiliary cells terminally on brief filaments which grow 

from the supporting cells of the carpogonial filaments before fertilization; cystocarps 

enclosed in cup- or vase-like pericarps; the thalli (cylindrical or flattened, branched 

or unbranched) usually hollow. Champia may be regarded as the standard genus. 

In various red algae, the germinating carpospore or tetraspore gives rise to a globe 
of cells which grows to produce the thallus (Kylin, 1917). In the present group the 
sporeling is particularly blastula-like. Its upper layer of cells becomes a ring of apical 
cells, of definite number, distinguishing the group from others which grow by apical 
cells either of a single filament or of a fascicle of indefinite number. The apical cells 
are indeed homologous with the apical cells of filaments, but the cells derived from 
them are arranged in a three-dimensional pattern as in the tissues of higher organisms; 
it is only in the reproductive structures that the filamentous structure remains evident. 
The order thus limited by Kylin (1932) is a specialized group including only the 
two families Rhodymeniacea [Rhodymeniaceae] Hauck and Champiea [Champieae] 
Kiitzing. The latter family is the more specialized; the hollow thalli are partitioned 
by transverse septa and the supporting cells produce usually just two auxiliary cells. 
In many examples of this family, after fertilization and the fusion of the zygote with 
the auxiliary cells, the latter proceed to unite with further neighboring cells to pro- 
duce a massive coenocyte from which the brief carpospore-bearing filaments arise. 
The resulting structure is deceptively similar to that which occurs in the Corallinea. 
The parasite Faucheocolax and its host Fauchea belong to this order. 

Order 6. Floridea [Florideae] C. Agardh Syst. Alg. xxxiii (1824). 

Order Floridees Lamouroux in Ann. Mus. Hist. Nat. Paris 20: 115 (1813). 

Section Florideae C. Agardh Synops. Alg. Scand. xiii (1817). 

Orders Trichoblasteae, Axonoblasteae, and Platynoblasteae Kiitzing Phyc. Gen. 

370,413,442 (1843). 
Orders Ceramieae, Spyridicae, Chondrieae, and Rhodomeleae J. Agardh Sp. 

Alg. vol. 2 (1851-1863). 
Ceramiales Oltmanns Morph. u. Biol. Alg. 1: 683 (1904). 
Order Ceramiales Kylin in Kgl. Svensk. Vetensk.-Akad. Handl. 63, no. 11 : 132 
The Floridees of Lamouroux included the whole group of red algae organized as 
four genera, Chondriis Stackhouse and the new genera Claudea, Delesseria, and 
Gelidium. Lamouroux listed first Claudea and Delesseria, belonging to the present 
order, to which the name is accordingly applied. 

This order is characterized by specialized strict patterns in the development of the 
feniale reproductive structures. The carpogonial filament is always of four cells. The 
supporting cell initiates, in definite patterns, brief additional filaments. After fertili- 
zation, the supporting cell cuts off one more cell adjacent to the zygote, and this be- 
comes the auxiliary cell. The spore-bearing structures developed from it are naked 
in the more primitive examples; in most, they are protected by pericarps, which, in 
some examples, begin to develop before fertilization. 

There are four families, all numerous in species: Ceramiea (Harvey) Kutzing, 

52 ] The Classification of Lower Organisms 

Dasyea Kiitzirxg, Delesseriea Kutzing, and Polysiphoniea Kiitzing [Rhodomelaceae 
Hauck). The Ceramiea are mostly filaments, uniseriate or becoming pluriseriate by 
lengthwise divisions. In many members of the other families the bodies are thallose, 
though consisting essentially of filaments produced in definite patterns. In many 
Delesseriea the branches of the thalli simulate leaves of higher plants. 

Gonimophyllum is parasitic on Botryoglossum; both are Delesseriea. Various 
species of Janczewskia, a genus of Polysiphoniea, attack Laurencia, Chondria, and 
other members of the same family. This was the first genus of parasitic red algae to 
be recognized as such, by Solms-Laubach (1877). 

Such are the red algae. The Bangialea appear to represent the transition between 
the organisms which lack nuclei and the generality of nucleate organisms. The 
Heterocarpea appear to be a specialized offshoot, leading to no other group. 

Chapter VI 

Phylum 2. PHAEOPHYTA Wettstein 

FucoroEAE C. Agardh Synops Alg. Scand. ix (1817). 

Orders Diatomeae and Fucoideae C. Agardh Syst. Alg. xii, xxxv (1824). 

Stamme Diatomea and Fucoideae Haeckel Gen. Morph. 2: xxv, xxxv (1866). 

Stamme Zygophyta in part and Phaeophyta Wettstein Handb. syst. Bot. 1: 71, 
171 (1901). 

Divisions Zygophyceae in part and Phaeophyceae Engler Syllab. ed. 3: 8, 15 

Chysophyta, with subordinate groups Chrysophyceae, Bacillariales, and Hetero- 
kontae, Pascher in Ber. deutschen bot. Gess. 32: 158 (1914). 

Stamm Chrysophyta Pascher in Siisswasserfl. Deutschland 11: 17 (1925). 

Phyla Chrysophycophyta and Phaeophycophyta Papenfuss in Bull. Torrey Bot. 
Club 73: 218 (1946). 

Organisms typically living by photosynthesis, without chromoprotein pigments, 
the plastids containing chlorophylls a and c, carotin, and various xanthophylls. Lutein 
(the xanthophyll of typical plants) may be present but is usually exceeded in quantity 
by flavoxanthin, violoxanthin, isofucoxanthin, or fucoxanthin, particularly the last. 
The xanthophylls occur usually in quantity sufficient to give the organisms a yellow 
or brown color. True starch is not produced. Many examples contain granules of a 
white solid called leucosin, presumably a carbohydrate, which does not give a blue 
color with iodine. The cells are usually enclosed in walls consisting of cellulose to- 
gether with larger quantities of other carbohydrates or oxidized or esterized carbo- 
hydrates. Silica or calcium carbonate may be deposited. Methanol extracts of the 
cells contain fucosterol, a sterol distinct from the sitosterol of typical plants. Flagel- 
late cells are usually produced; these bear one pantoneme or pantacroneme flagellum, 
and usually, in addition, one acroneme or simple flagellum. Exceptional examples, 
non-pigmented or without flagellate stages, are rather numerous. The obvious stand- 
ard genus of the phylum is Fucus L. 

The chemical characters are stated on the authority chiefly of Carter, Heilbron, 
and Lythgoe (1939), Miwa (1940), and Tseng (1945). The character of the flagel- 
lation, positively known of rather few examples, is stated by authority of Petersen 
(1929), Vlk (1931, 1939), Couch (1938, 1941), Longest [1946), Manton (1952), 
and Ferris (1954). 

These characters bind together an assemblage of organisms which is in some re- 
spects original herel. Engler (1897), West (1904), and Smith (1918, 1920) included 
the chrysomonad flagellates in the group of brown algae. Pascher (1914) combined 
as or!e group the chrysomonads, the diatoms, and the exceptional green algae called 
Heierokontae. Later (1927, 1930), he included also the colorless flagellates of family 
Moiiadina. He did not associate this group with the brown algae, and subsequent 
authors have in general followed him. Kylin ( 1933 ) , however, considered the diatoms 
to be the closest allies of the brown algae, both groups being descended from the 
brown flagellates. Almost certainly, he was correct. Couch showed that the paired 
unlike flagella of the typical Oomycetes are respectively pantoneme and acroneme, 

iManton (1952) recognized this group, but omitted nomenclatural formalities. 


The Classification of Lower Organisms 

Fig. 8. — Ochromonadalea: a, b, Chrysocapsa paludosa after West (1904); a, a 
colony; b, zoospores. C-f, Phaeocystis globosa after ScherlTel (1900); c, a colony 
X 50; d, a cell with two plastids, a mass of leucosin forming on a mound of proto- 
plasm projecting into the central vacuole; e, production of zoospores; f, a zoospore. 
g, h. Cell and statospore of Ochromonas granularis after Doflein (1922). i, Cell of 
Monas sp. j. Two cells of Brehmiella Chrysohydra after Pascher ( 1928) . k, A very 
young colony of Dendromonas virgaria after Stein (1878). 1, Colony of Ccphalo- 
thamnium Cyclopum after Stein, op. cit. m. Cells of Epipyxis utriculus after Stein, 
op. cit. n. Colony of Synura Uvella. x 1,000 except as noted. 

Phylum Phaeophyta [ 55 

and distinguished these fungi from practically all others by the presence of cellulose 
in their walls. 

The phylum thus assembled may be organized as four classes. 
1. Miscellaneous groups, mostly small and rela- 
atively unspecialized, of varied body type; not 

of the characters of the following groups Class 1. Heterokonta. 

1. Comparatively numerous and specialized 

2. Unicellular brown organisms with shells 

of silica consisting of two parts Class 2. Bacillariacea. 

2. Organisms of fungal or chytrid body 
types producing swimming spores with 

paired unlike fiagella Class 3. Oomycetes. 

2. Filamentous and thallose brown algae Class 4. Melanophycea. 

Class 1 . HETEROKONTA Luther 

Class Flagellata or Mastigophora Auctt., in part. 

Class Heterokontae Luther in Bihang Svensk. Vetensk.-Akad. Handl. 24, part 

3, no. 13: 19 (1899). 
Subclass Chrysomonadineae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, 

Abt. la: iv (1900). 
Class Silicoflagellatae (Borgert) Lemmermann in Ber. deutschen bot. Gess. 19: 

254 (1901). 
Phylum Siphonophyceae and class Vaucherioideae Bessey in Univ. Nebraska 

Studies?: 285, 286 (1907). 
Chrysophyceae and Heterokontae Pascher in Ber. deutschen bot. Gess. 32: 158 

Divisions Chrysophyceae and Heterokontae, and classes Chrysomonadineae , Rhizo- 

chrysidineae, Chrysocapsineae, Chrysosphaerineae, Chrysotrichineae, Hetero- 

chloridineae, Rhizochloridineae, Heterocapsineae, Heterococcineae, Hetero- 

trichineae, and Heterosiphoneae Pascher in Beih. bot. Centralbl. 42, Abt. 2: 

323,324 (1931). 
Classes Ebriaceae, Silico flagellata, and Coccolithophoridae Deflandre, and Chrys- 

omonarfina Hollande in Grasse Traite Zool. 1, fasc. 1: 407,425,438, 471 (1952). 
Class Phytomastigophorea Hall Protozoology 117 (1953), in part. 
Phaeophyta which lack the distinctive characters of the remaining three classes. 
Luther named the group on the occasion of his discovery of Chlorosaccus, and this 
genus may be regarded as the type. 

The chrysomonad flagellates are the core of this class and of the first two among 
the five orders into which it is divided. In the classification of these two orders, three 
novelties will be noted. 

(a) Pascher (1913) made of the chrysomonad flagellates three orders character- 
ized respectively by paired unequal flageila, paired equal flagella, and solitary 
fiagella. Petersen (1929) found that the supposedly equal fiagella of Synura are 
actually unlike, being respectively pantoneme and acroneme. Here, accordingly, 
Pascher's first two orders are combined. 

(b) Pascher made separate classes or orders of groups related to the chrysomonad 
flagellates but of distinct body type, as palmelloid, chlorococcoid, filamentous, or 


The Classification of Lower Organisms 

Fig. 9. — Ochromonadalea : a, Mallomonas roseola, based on Stein (1878) and 
Conrad (1926). h, Syracosphaera Quadricornu; c, Calyptosphaera insignis; d, Cal- 
ciconus vitreus; after Schiller (1925). Silicoflagei.lata: e, f, Colony and zoospore 
of Epichrysis after Pascher (1925). g, Part of the thallose growth of Hydrurus 
foetidus. h, Cell, and i, j, statosporos of Chromulina Pascheri after Hofeneder 
(1913). k, 1, Skeletons of Dictyocha Fibula and Distephanus Speculum from di- 
atomaceous earth at Lompoc, California, m, Rhizochrysis Scherffeli after Doflein 
(1916). Mix 1,000. 

Phylum Phaeophyta [57 

amoeboid. By Pascher's own principle of the repeated evolution of body types, 
these groups are surely artificial. Here most of them are broken up and their mem- 
bers distributed between the two chrysomonad orders according to whether the 
flagella of their motile stages are paired or single. It is not possible to divide by this 
character ameboid forms not known to produce flagellate stages; these are lumped 
in the second order. 

fc) Since flagella appear to have evolved as a device for the dissemination of 
unicellular pigmented organisms, examples whose vegetative state is that of clusters 
of non-motile cells are placed in each order before those which are flagellate in 
the vegetative condition. 

The two chrysomonad orders are particularly characterized by production of 
leucosin. They are further characterized by production of resting cells of a type 
called statospores. This occurs by the deposition within the protoplast of a globular 
shell impregnated with silica, punctured by a single pore, and often marked on the 
outer surface by warts, spines, or ridges, of definite pattern. The external protoplasm 
migrates through the pore to the interior of the shell, and the pore is then closed 
by deposition of a silicified plug. 

The group which is treated as the third order of the present class includes the 
typical Heterokonta. Compared with typical green algae, these organisms give the 
impression of a markedly distinct class; placed next to the chrysomonads, they 
appear scarcely entitled to this rank. Their name is the oldest applicable to the 
present class, and is accordingly so applied. If it appear expedient to maintain the 
typical Heterokonta as a distinct class, the remainder of the present one will be 
called Chrysomonadinea [Chrysomonadineae] (Engler) Pascher. 

Of including the choanoflagellates and anisochytrids in the present class as addi- 
tional orders, one may say that it is not contrary to current knowledge. 
1. Mostly pigmented; non-pigmented examples 
mostly producing motile cells with two 

2. Brown or colorless. 

3. Producing motile cells with two 

flagella (exceptionally more) Order 1. Ochromonadalea. 

3. Producing motile cells with one 
flagellum; or without known flagel- 
late stages Order 2. Silicoflagellata. 

2. Green Order 3. Vaucheriacea. 

1. Non-pigmented, producing motile cells with 
one flagellum. 

2. Predatory, flagellate in the vegetative 
condition, each cell bearing a collar-like 

protoplasmic ridge Order 4. Choanoflagellata. 

2. Parasitic or saprophytic, the vegetative 

cells non-motile, walled Order 5. Hyphochytrialea. 

Order 1. Ochromonadalea [Ochromonadales] Pascher Siisswasserfl. Deutschland 

2: 10,51 (1913). 
Suborder Monadina Biitschli in Bronn KI. u. Ord. Thierreichs 1 : 810 (1884). 
Order Isochrysidales Pascher op. cit. 10, 42. 
Order Syracosphaerinae Schiller in Arch. Prot. 51: 108 (1925). 

58] The Classification of Lower Organisms 

Orders Heliolithae and Orthlithinae Deflandre in Grasse Traite Zool. 1, fasc. 
1: 452, 457 (1952). 
Brown or colorless Heterokonta, the swimming cells of typical examples with 
two flagella which are respectively pantoneme and acroneme. In the exceptional 
family Trimastigida there are a pair of equal flagella and a third flagellum shorter 
or longer than these; the detailed structure of the flagella of this family is unknown. 
Cells of pigmented types contain usually one or two lateral band-shaped plastids. 
Details of nuclear division are known chiefly by the observations of Doflein (1918, 
1922) on Ochromonas. The flagella spring from a granule which may be identified as 
a blepharoplast, near which lies the nucleus. The blepharoplast is connected through 
two stainable strands (rhizoplasts) to two granules, recognizable as centrosomes, on 
the two sides of the nucleus. The spindle forms within the intact nuclear membrane 
with its poles at the centrosomes. The chromosome number appears to be about 4. 
The nuclear membrane presently disappears. At metaphase, the rhizoplasts are found 
to lead to separate blepharoplasts, each bearing two flagella. Sexual processes are 
scarcely known in this group. Schiller (1926) observed in Dinobryon the division of 
calls into two which are released to swim and conjugate in pairs. 

This order is believed to represent the direct ancestry of the two following, and 
also of the typical brown algae. 
1. Not filamentous. 

2. Flagellate stages with a pair of equal 
flagella and a third which is shorter or 

longer Family 1. Trimastigida. 

2. Flagellate stages with two unequal 

3. Without calcareous structures at- 
tached to the cell walls. 

4. Cells not enclosed in loricae, 
i. e., open shells. 

5. Not flagellate in the vege- 
tative condition Family 2. Chrysocapsacea. 

5. Flagellate in the vegeta- 
tive condition, not forming 
free-swimming circular or 

globular colonies Family 3. Monadina. 

5. Free-swimming circular or 

globular colonies Family 4. Syncryptida. 

4. Cells enclosed in loricae Family 5. Dinobryina. 

3. With calcareous structures at- 
tached to the cell walls Family 6. Hymenomonadacea. 

1. Filamentous Family 7. Phaeothamnionacea. 

Family 1. Trimastigida [Trimastigidae] Kent Man. Inf. 1: 307 (1880). Family 
Trimastigaceae Senn in Engler and Prantl. Nat. Pflanzcnfam. I Teil, .\bt. la: 141 
(1900). Family Prymncsiidae Hall Protozoology 127 (1953). Organisms producing 
swimming cells with a pair of equal flagella and a third flagellum longer or shorter 
than these. With a vegetative stage as globular non-motile colonies as large as pin- 
heads, of pigmented cells; marine: Phacocystis. Motile solitary cells, pigmented: 
Prymncsium, Chrysochromidina; Platychrysis with an amoeboid stage. Motile 
solitary cells, not pigmented: Dallingeria, Trimastix, Macromastix. 

Phylum Phaeophyta [ 59 

Family 2. Chrysocapsacea [Chrysocapsaceae] Pascher in Siisswasserfl. Deutschland 
2: 85 (1913). Family Chrysocapsidae Poche in Arch. Prot. 30: 156 (1913). Non- 
motile cells with brown plastids (usually two), imbedded in gelatinous matter and 
forming colonial aggregates, the protoplasts sometimes escaping as zoospores with 
two flagella. Chrysocapsa Pascher, in fresh water, the colonies few-celled. Phaeo- 
sphaera West and West, the colonies more extensive. 

Family 3. Monadina Ehrenberg Infusionsthierrhen 1 (1838). Family Monades 
Goldfuss ( 1818), the mere plural of a generic name. Family Dendromonadina Stein 
Org. Inf. 3, I Halfte: x (1878). Family Monadidae Kent (1880). Family Hetero- 
monadina Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 815 (1884). Family Chryso- 
monadaceae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 2: 570 
(1897), not family Chrysomonadina Stein. Family Ochromonadaceae Senn in 
Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. la: 163 (1900). Family Ochro- 
monadidae Doflein. Pigmented or colorless Ochromonadalea, flagellate in the 
vegetative condition, not forming circular or globular free-swimming colonies, nor 
loricate, nor bearing calcareous structures on the cell walls (these being the distinc- 
tions respectively of the three following families). 

Ochromonas is considered to be in its normal condition when it occurs as solitary 
swimming cells; it occurs also as gelatinous colonies like those of Chrysocapsa. 
Stylochrysalis consists of O chromonas-like cells attached by a stalk at the end away 
from the flagella. Chrysodendron is similar but colonial, the cells attached by branched 
stalks. Brehmiella Pascher (1928) may occur as free-swimming Ochromonas-Vikt 
cells, or these may become attached by the end away from tlie flagella and develop 
a whorl of pseudopodia at the free end. Pseudopodia are a device for predatory nutri- 
tion, here occurring in an organism which is capable also of photosynthesis. Hetero- 
chromonas includes organisms of the structure of Ochromonas but without plastids, be- 
ing presumably saprophytic, and containing only a pigmented speck by which it is sup- 
posed that the direction of light is perceived. The historical generic name Monas 
O. F. Miiller, as restricted in application by scholars up to Ehrenberg and as applied 
ever since, designates totally non-pigmented cells, saprophytic or predatory, free- 
swimming like Ochromonas or attached like Stylochrysalis [Physomonas Kent desig- 
nates cells of Monas in the attached condition). There are believed to be several 
species, but the group remains poorly known. It was in some member of it that Loeffler 
(1889) first observed the pantoneme character of flagella. Dendromonas consists of 
similar cells forming colonies like those of Chrysodendron. In Cephalothamnium 
Stein, Monas-\ikc cells are gathered in capitate clusters on stout stalks. Anthophysis 
Bory is an organism which Leeuwenhoeck had described as a microscopic water 
plant: it consists of Monas-Vikt cells at the ends of branching stalks colored yellow 
by deposits of iron. The comparatively unfamiliar original spellings of the two 
generic names just mentioned were restored by Kudo ( 1946). The name Uvella Bory 
appears to represent small clusters of cells of Cephalothamnium or Anthophysis 
which have broken loose to swim free. 

Family 3. Syncryptida [Syncryptidae] Poche in Arch. Prot. 30: 156 (1913). Family 
Isochrysidaceae Pascher in Siisswasserfl. Deutschland 2: 43 (1913), not based on a 
generic name. Family Isochrysidae Calkins Biol. Prot. 262 (1926). Families Synura- 
ceae and Syncryptaceae Smith Freshw. Algae (1933). Ochromonas-Vike cells forming 
circular or globular free-swimming colonies. Flagella markedly unequal, colonies 
circular: Cyclonexis; colonies globular: Uroglena, Uroglenopsis. Flagella apparently 
equal: Syncrypta, Synura. 

60 ] The Classification of Lower Organisms 

Family 4. Dinobryina Ehrenberg Infusionsthierchen 122 (1838). Family Dino- 
hryaceae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 2: 570 (1897). 
Pigmented or colorless cells of the characters of Ochromonas or Monas, sheltered in 
loricae, that is, in transparent open shells, solitary or colonial. The pigmented 
examples have generally been referred to Ochromonadaceae (or whatever), the 
colorless to Monadidae (or whatever). Pigmented, solitary, flagella markedly 
unequal: Epipyxis, Stylo pyxis; flagella apparently equal: Chry so pyxis Stein {Dere- 
pyxis Stokes). Pigmented, forming branching colonies: Dinobryon, Hyalobryon. 
Poteriochromonas Scherffel resembles Stylopyxis, but the protoplast can project 
pseudopodia from its lorica, thus supplementing photosynthesis by predatory nutri- 
tion. Non-pigmented, solitary, flagella markedly unequal: Stokesiella; flagella ap- 
prrently equal: Diplomita. Non-pigmented cells in colonies quite of the character 
of those of Dinobryon: Stylobryon. 

Family 5. Hymenomonadacea [Hymenomonadaceae] Senn in Engler and Prantl 
Nat. Pflanzenfam. I Teil, Abt. la: 159 (1900). Family Coccolithophoridae Lohman 
in Arch. Prot. 1: 127 (1902). Family Hymenomonadidae Doflein. Family Cocco- 
lithidae Poche in Arch. Prot. 30: 157 (1913). Order Syracosphaerinae and family 
Pontosphaeraceae Schiller in Arch. Prot. 51: 8 (1925). Families Syracosphaeraceae, 
Halopappaceae, Deutschlandiaceae, and Coccolithaceae Kampter. Family Thora- 
cosphaeracee Schiller in Rabenhorst Kryptog.-Fl. Deutschland ed. 2, 10, Abt. 2: 156 
(1930). Y3imi\it5 Syracosphaeridae, Calcisolenidae, Thoracosphaeridae, and Braad- 
rudosphaeridae Deflandre in Grasse Trait6 Zool. 1, fasc. 1: 452, 457, 458 (1952). 
Family Discoasteridae Tan Sin Hok. Suborder Coccolithina Hall Protozoology 130 
(1953). Solitary cells with one or two brown plastids, usually with two apparently 
equal flagella, having a thin cell wall from which project bodies of calcium carbonate 
(coccoliths) of definite form. 

More than twenty genera and nearly 150 species have been described (Lohman; 
Schiller; Kamptner, 1940). Neither the number of species nor the variety of form 
appears to warrant making more than one family of the group. Nearly all examples 
are marine. In Pontosphaera, Calyptosphaera, and allied genera, the coccoliths are 
disks or hemispheres, sometimes umbonate and sometimes marked by one or more 
pits. In Syracosphaera the coccoliths, or a few of them near the insertion of the 
flagella, bear horn-like projections. In Najadea, Halopappus, and Calciconus, each 
cell bears a whorl or elongate bristles. Cells of Calcisolenia are fusiform, without 
flagella, with an armor of two layers of spiral bands of calcareous matter. In Hymen- 
omonas and Coccolithus Swartz 1894 [Coccosphaera Wallich 1877, non Perty 1852; 
Coccolithophora Lohman 1902) the coccoliths are punctured and accordingly ring- 
shaped; Hymenomonas difi"ers from most of the group in occurring in fresh water. In 
Discosphaera and Rhabdosphaera the punctured calcareous bodies are drawn out to 
the form of tubes, spools, or trumpets. 

These obscure organisms are not without importance. They occur in all oceans, 
being most abundant in gulfs, such as the Adriatic, where the salinity is diminished 
by rivers (Schiller, 1925). According to Bernard (1947) turbidity in the Mediter- 
ranean depends chiefly on this group. Coccoliths are abundant in the ooze on the 
bottoms of oceans. They occur as fossils as far back as the Cambrian, being par- 
ticularly abundant in certain Cretaceous deposits. 

Family 6. Phaeothamnionacea [Phaeothamnionaccae] Pascher in Siisswasserfl. 
Deutschland2: 113 ( 1913). Family Chrysotrichaceae Fascher (1914). Family Nema- 
tochrysidaceae Pascher (1925). Brown organisms, minute, marine, epiphytic, filamen- 

Phylum Phaeophyta [ 61 

tous, reproducing by zoospores bearing paired unequal flagella. Nematochrysis, the 
filaments unbranched; Phaeothamnio7i, the filaments branched. These organisms are 
believed to represent the transition between the Chrysocapsacea and the typical 
brown algae. 

There is a family Amphimonadidae or Amphimonadaceae of unwalled colorless 
flagellates with paired supposedly equal flagella. They appear to belong to the 
kingdom of plants, in the neighborhood of Chlamydomonas and Polytoma. If, how- 
ever, future study shows their flagella actually to be respectively pantoneme and 
acroneme, they are to be placed in the present order. 

Order 2. Silicoflagellata Borgert in Zeit. wiss. Zool. 51: 661 (1891). 
Chromomonadina Klebs in Zeit. wiss. Zool. 55: 394 (1893). 
Order Chromomonadina Blochmann Mikr. Tierwelt ed. 2. Abt. I: 57 (1895). 
Subclass Chrysomonadineae Engler in Engler and Prantl Nat. Pflanzenfam. 

ITeil, Abt.'la: iv (1900). 
Order Chrysomonadales Engler Syllab. ed. 3: 7 (1903). 

Chrysomonadinae; Euchrysomonadinae , with order Chromulinales; Chryso- 
capsinae; and Rhizochrysidinae Pascher in Siisswasserfl. Deutschland Heft 2 
Chrysomonadales, Chrysocapsales, Chrysosphaerales, and Chrysotrichales Pas- 
cher in Ber. deutschen bot. Gess. 32: 158 (1914). 
Order Chrysomonadina Doflein Lehrb. Prot. ed. 4: 401 (1916). 
Order Chrysomonadida Calkins Biol. Prot. 258 (1926). 

Classes Chrysomonadineae , Rhizochrysidineae, Chrysocapsineae, Chrysosphaeri- 
neae, and Chrysotrichineae Pascher in Beih. bot. Centralbl. 48, Abt. 2: 323 
Suborders Euchrysomonadina, Silicoflagellina, Rhizochrysidina, and Chrysocap- 
sina Hall Protozoology 125, 128, 130, 132 (1953). 
Organisms of much the character of Ochromonadalea, but producing flagellate 
stages with a single flagellum, or not producing flagellate stages. The detailed 
structure of the flagella has seemingly never been determined. Statospores are known 
to be produced by Chromulina, Mallonionas, and (of somewhat exceptional charac- 
ter) by Hy drums. Sexual reproduction has not been observed. Mitosis, with an 
intranuclear spindle and numerous chromosomes, was observed by Doflein (1916) 
in Rhizochrysis. 

This order is supposed to represent the direct ancestry of orders Choanoflagellata 
and Hyphochytrialea. 

1. Neither amoeboid nor truly filamentous. 
2. Not flagellate in the vegetative condi- 

3. Microscopic colonies Family 1. Chrysosphaeracea. 

3. Macroscopic gelatinous colonies 

simulating filaments Family 2. Hydruragea. 

2. Flagellate in the vegetative condition. 
3. Without prominent siliceous struc- 
tures Family 3. Chrysomonadina. 

3. With siliceous scales usually bearing 

bristles Family 4. Mallomonadinea. 

3. With siliceous internal skeletons Family 5. Actiniscea. 

62 ] The Classification of Lower Organisms 

1. Amoeboid Family 6. Chrysamoebida. 

1. Filamentous Family 7. Thallochrysidacea. 

Family 1. Chrysosphaeracea [Chr>'Sosphaeraceae] Pascher in Arch. Prot. 52: 562 
(1925). Family Naegelliellaceae Pascher op. cit. 561. Family Nagelliellidae Hall 
Protozoology 133 (1953). Non-motile brown cells, either capable of repeated division 
into two, thus forming aggregates of indefinite number, or else undergoing multiple 
division and producing colonies of definite number of cells; mostly known to produce 
uniflagellate zoospores. Chrysosphaera, Epichrysis, Chrysospora, Gloeochrysis, Nae- 
gelliella, and other genera. 

Family 2. Hydruracea [Hydruraceae] West British Freshw. Algae 45 (1904). 
Hydrurina Klebs in Zeit. wiss. Zool. 55: 420 (1893). Family Hydruridae Poche in 
Arch. Prot. 30: 158 (1913). Like Chrysosphaeracea, but the colonies dendroid, 
growing at the tips, becoming macroscopic; producing tetrahedral zoospores and 
spheroidal resting cells bearing a unilateral crest. Hydrurus foetidus, in mountain 

Family 3. Chrysomonadina Stein Org. Inf. 3, I Halfte: x (1878). Family 
Chrysomonadidac Kent Man. Inf. (1880). Family Chromulinaceae Engler in Engler 
and Prantl Nat. Pflanzenfam. I Teil, Abt. 2: 570 (1897). Family Chromulinidae 
Doflein. Brown flagellates with a single anterior flagellum, sometimes producing 
siliceous granules but without more extensive siliceous structures. Free-swimming, 
walled: Chrysococcus, Microglena. Naked: Chromulina, the type genus of Chryso- 
monadina, the generic name Chrysomonas being a synonym. Organisms of this genus 
are rather freely capable of producing pseudopodia and supplementing photosynthetic 
nutrition by predatism, or, alternatively, of producing gelatinous aggregates of 
walled non-motile cells (Hofender, 1913; Gicklhom, 1922). Chrysapsis differs from 
Chromulina in having in each cell a single plastid in the form of a network. Solitary 
attached cells, producing pseudopodia only occasionally: Lepo chromulina. Bearing 
whorls of permanent pseudopodia: Cyrtophora, Pedinella, Palatinella (Pascher, 

Family 4. Mallomonadinea Diesing in Sitzber. Akad. Wiss. Wien Math.-Nat. CI. 
52, Abt. 1: 304 (1866). Family Mallomonadidae Kent (1880). Brown uniflagellate 
free-swimming cells with an armor of siliceous scales usually bearing bristles. Mallo- 
m.onas, solitary cells, the bristle-bearing scales circular. Conradiella, the scales of the 
form of rings about the body. Chrysosphaerella, spherical colonies, each cell with two 
long bristles. 

Family 5. Actiniscea [Actinisceae] Kiitzing Phyc. Germ 117 (1845). Family 
Dictyochidae Wallich. Class Silicoflagellata (Borgert), orders Siphonotestales and 
Stereotestales, and families Dictyochaceae and Ebriaccae Lemmermann in Ber. 
deutschen bot. Gcss. 19: 254-268 ( 1901 ). Division (?) Silicoflagellatac Engler. Family 
SiHcoflagellidae Calkins Biol. Prot. 263 (1926). Famihes Ebriopsidae, Ditripodiidae, 
Ammodochidae, and Ebriidae Deflandre in Grasse Traite Zool. 1, fasc. 1: 421, 423, 
424 (1952). Solitary brown uniflagellate cells with a continuous internal skeleton of 
silica. Marine, commonest in colder oceans. 

The skeletons are not subject to decay and are found as micro fossils in chalk 
and diatomaceous earth. They have been reported from the Silurian and are com- 
monest in certain Cretaceous deposits. Ehrcnbcrg described several fossil species, 
classifying them as diatoms. The living forms, subsequently discovered, include 
apparently the same species. 

Gemeinhardt (in Rabcnhorst, 1930) accounted for the structure of the cells. 

Phylum Phaeophyta [ 63 

They are approximately of radial symmetry, the axis being shorter than the diameter. 
The skeleton is completely imbedded in protoplasm. It may be a mere ring; or the 
ring may bear radially projecting spines; or it may be the margin of a more or less 
complicated basket-shaped network coaxial with the cell. Numerous brown plastids 
lie near the surface of the protoplast. There is no cell wall. The double cells, like 
two cells lying face to face, which have occasionally been seen, are not stages of 
conjugation, but of cell division, in which one daughter cell retains the original 
skeleton while the other develops a new skeleton in the position of a mirror image 
of the original one. 

Lemmermann and Gemeinhardt accounted for only six genera and twenty-four 
species, but Gemeinhardt recognized numerous varieties, and it is probable that the 
number of species has been underestimated. Mesocaena, the skeleton a mere ring, 
smooth or spiny; Dictyocha, Distephanus, Cannopilus, the skeleton more or less 

Family 6. Chrysamoebida [Chrysamoebidae] Poche in Arch. Prot. 30: 157 (1913). 
Families Rhizochrysidaceae , Chrysarachniaceae, and Myxochrysidaceae Pascher in 
Beih. Bot. Centralbl. 48, Abt. 2: 323 (1931). Family Rhizochrysididae Hollande in 
Grasse Traite Zool. 1, fasc. 1: 547 (1952). Families Rhizochrysidae and Myxochry- 
sidae Hall Protozoology 130, 132 (1953). Amoeboid organisms with brown plastids 
of the form of one or two parietal films in each cell. Rhizaster, an attached organism 
resembling Cyrtophora and Pedinella but lacking the flagellum. Chrysocrinus, at- 
tached to algae, the protoplast covered by a dome-shaped shell punctured by many 
pores through which project the slender psudopodia. Chrysamoeba, a freely moving 
cell usually with one flagellum; Rhizochrysis, similar, without the flagellum. Myxo- 
chrysis, a large multinucleate form. Chrysarachnion, the cells clustered and linked 
together by strands of protoplasm. Lagynion, having an attached vase-shaped lorica 
from which projects usually a single slender pseudopodium. Chrysothylakion, with 
a retort-shaped lorica from which project many slender pseudopodia, branching 
and anastomosing. Only the plastids distinguish these organisms from various genera 
classified as Rhizopoda, Heliozoa, or Sarkodina. 

Family 7. Thallochrysidacea [Thallochrysidaceae] Pascher (1925). Brown or- 
ganisms producing definite filaments of walled cells and reproducing by anteriorly 
uniflagellate zoospores. T hall ochry sis. Phaeodermatium. 

Order 3. Vaucheriacea [Vaucheriaceae] Nageli Gatt. einzell. Alg. 40 (1849). 

Class Heterokontae and orders Chloromonadales and Confervales Luther in 
Bihang Svensk. Vetensk.-Akad. Handl. 24, part 3, no. 13: 19 (1899). Not 
Chloromonadina Klebs (1893); not order Confervoidea C. Agardh (1824). 

Vaucheriales Bohlin Grona Algernas 25 (1901). 

Order Vaucheriales Clements Gen. Fung. 14 (1909). 

Orders Heterochloridales, Heterocapsales, Heterococcales, Heterotrichales, and 
H eter osiphonales mlitdwigizbZ: 10-21 (1912). 

Division Heterokontae, Classes Heterochloridineae, Rhizochloridineae, Hetero- 
capsineae, Heterococcineae, Heterotrichineae , and Heterosiphoneae, and or- 
ders Rhizochloridales and Botrydiales Pascher in Beih. bot. Centralbl. 48, 
Abt. 2: 324 (1931). 

Class Xanthomonadina with orders Heterochloridea and Rhizo chloride a De- 
flandre in Grasse Traite Zool. 1, fasc. 1: 212, 217, 220 (1952). 

Order Heterochlorida Hall Protozoology 133 (1953). 


The Classification of Lower Organisms 

Organisms producing motile cells with paired unequal flagella which Vlk (1931) 
found to be respectively pantoneme and acroneme, differing from Ochromonadalea 
in being of a green or yellow-green color, and in being mostly of algal body type, 
i. e., walled and non-motile. The cell wall consists usually of two parts which become 
separate when the cell divides; the two parts are believed to be distantly homologous 
with the wall and pRig of the statospores of Ochromonadalea and Silicoflagellata 
(Pascher, 1932). The storage products are oil and sometimes leucosin. 

As this is the group to which the class name Heterokontae was first applied, it is 

f-'^y-'v'^^^; ■■■• >i%'X°-^'^^ 

Fig. 10. — Vaucheriacea: a, b^ Chlorosaccus fluidus, cells of the colony and zoo- 
spores, after Luther (1899). c, d^ Chlorarnoeba heteromorpha x 1,000 after Bohlin 
(1897). e, f, g. Cell, empty cell, and zoospores of Characiopsis gibba x 1,000 after 
Pascher (1912). h, Dioxys Incus after Pascher (1932). i, j, k, Cell, edge of cell, 
and statospore of Pseudotetraedron neglectum x 1,000 after Pascher (1912). 1, Spi- 
rodiscus fulvus x 1,000. m. End of a filament of Tribonema bombycina x 1,000. 
n, Antheridium and oogonia of Vaucheria Gardneri x 100. o. Filament of Vaucheria 
sessilis x 100. 

Phylum Phaeophyta [ 65 

the type group of the class. As established by Luther, the class consisted of the new 
genus Chlorosaccus together with a few genera of flagellates ( Vacuolaria was included 
in error) and a few transferred from the group of typical green algae. From time to 
time, other green algae have been transferred, and it has become evident that the 
group is a fairly extensive one. Green organisms can be recognized as belonging here 
by a negative reaction to the iodine test for starch, and by the fact that they give a 
b'uish color when heated with hydrochloric acid, instead of a yellow one, as typical 
green algae do: the difference depends upon differences in the complement of 
photosynthetic pigments. Bohlin (1901) placed Vaucheria here; most authors have 
not followed him, but Smith (1950) has done so. This genus brings with itself the 
oldest name for the group as an order. 

Mitosis in Vaucheria was described by Hanatschek (1932) and Gross (1937). The 
spindle is intranuclear; Hanatschek saw centrosomes at the poles. The conjugation of 
equal free-swimming gametes was observed in Tribonema and several other genera by 
ScherfFel (1901), and in Botrydium by Rosenberg (1930). Vaucheria was one of the 
organisms by study of which the nature of fertilization was discovered (Pringsheim, 
1855). Hanatschek and Gross found that the first two divisions of the nucleus of 
the zygote are meiotic: the soma is haploid. 

This order is believed to represent the direct ancestry of the two following classes, 
Bacillariacea and Oomycetes. 

Pascher (1912, 1925) arranged the green Heterokonta in subordinate groups 
parallel to those of the typical green algae; and, as the main groups of green algae 
are treated as orders, he treated these groups also as orders (in 1931 as classes). 
They are scarcely entitled to such rank: too many of the classes or orders are of 
single families, and too many of the families are of one or two genera. Here, then, 
Pascher's classes and orders are suppressed and several of his families are reduced. 
1. Not truly filamentous nor producing rhizoids. 
2. The cells walled. 

3. Cells regularly dividing into two, 
forming gelatinous colonies; occa- 
sionally producing small numbers 
of zoospores. 

4. The colonies globular or Irreg- 
ular, becoming macroscopic Family 1. Chlorosagcacea. 

4. The colonies dendroid, micro- 
scopic Family 2. Mischococgacea. 

3. Cells normally undergoing division 
into several. 

4. Producing zoospores Family 3. Chlorotheciacea. 

4. Producing no motile cells Family 4. Botryococcagea. 

2. The cells loricate Family 5. Stipitogogcacea. 

2. The cells amoeboid Family 6. Chloramoebacea. 

1. Filaments of uninucleate cells Family 7. Tribonematagea. 

1. Cells becoming highly multinucleate, form- 
ing filaments or at least producing rhizoids Family 8. Phyllosiphonacea. 

Family 1. Chlorosaccacea [Chlorosaccaceae] Smith Freshw. Algae 145 (1933). 
Family Heterocapsaceae Pascher in Hedwigia 53: 13 (1912); there is no correspond- 
ing generic name. Gelatinous aggregates of cells which may divide, causing the 

66] The Classification of Lower Organisms 

aggregate to grow to macroscopic dimensions; or may produce one, two, or four 
zoospores. Chlorosaccus Luther, the standard genus of class Heterokonta. 

Family 2. Mischococcacea [Mischococcaceae] Pascher in Hedwigia 53 : 14 ( 1912). 
Microscopic colonies of globular cells joined by dichotomously branching gelatinous 
strands. Mischococcus. 

Family 3. Chlorotheciacea [Chlorotheciaceae] Luther in Bihang Svensk. Vetensk- 
Akad. Handl. 24, part 3, no. 13: 19 (1899). Families Chlorobotrydiaceae and Sci- 
adiaccae Pascher in Hedwigia 53: 17 (1912). Family Halosphaeraceae Pascher 
(1925). Family Ophiocytiaceae Auctt. Cells solitary, free or attached, capable of 
reproduction by division to form multiple zoospores, in some examples capable 
alternatively of producing multiple minute non-motile cells of the same form as 
the parent. Large free multinucleate cells, more or less globular: Botrydiopsis, Leu- 
venia. Smaller cells, elongate, curved or coiled: Characiopsis, Spirodiscus. Spirodiscus 
fuluus Ehrenberg in Abh. Akad. Wiss. Berlin 1830: 65 (1832) {nomen nudum) and 
Infusionsthierchen 86 (1838), whose identity has been a standing puzzle to bac- 
teriological systematists, is an older name of Ophiocytium parvidum (Perty) A. 
Braun (Copeland, 1954). It antedates the generic name Ophiocytium Nageli (1849); 
new combinations are required for the dozen additional species of this genus. The 
cells attached: some species of Characiopsis; Perionella; Dioxys. 

Family 4. Botryococcacea [Botryococcaceae] Pascher in Hedwigia 53: 13 (1912). 
Solitary or colonial cells reproducing strictly by production of non-motile cells. 
Botryococcus. Pseudotetraedron. 

Family 5. Stipitococcacea [Stipitococcaceae] Pascher in Beih. bot. Centralbl. 48, 
Abt. 2: 324 (1931). Family Stipitochioridae Deflandre in Grasse Trate Zool. 1, fasc. 
1: 221 (1952). Amoeboid cells with green plastids, partially enclosed in loricae at- 
tached to objects in water. Stipitococcus. 

Family 6. Chloramoebacea [Chloramoebaceae] Luther in Bihang Svensk. Vetensk.- 
Akad. Handl. 24, part 3, no. 13: 19 (1899). Family Chloramoebidae Poche in Arch. 
Prot. 30: 155 (1913). Families Heterochloridaceae and Rhizochloridaceae Pascher 
Siisswasserfl. Deutschland 11: 22, 26 (1925). Y^.miWts Heterochloridae, Rhizochlori- 
dae, Chlorarachnidae and Myxochloridae Deflandre in Grasse Traite Zool. 1, fasc. 1 : 
217-222 (1952). Amoeboid organisms with green plastids, without loricae, some- 
times swimming by means of paired unequal flagella. Chloramoeba, Chlorochromo- 
nas, Rhizochloris. 

Family 7. Tribonematacea [Tribonemataceae] Pascher in Hedwigia 53 : 19 ( 1912) . 
Family Confervaceae Luther (1899). Family Monociliaceae Smith Freshw. Algae 
160 (1933). Green Heterokonta producing filaments of uninucleate cells. The Lin- 
naean genus Conferva included a great variety of growths in water. Definite groups 
were separated from it, one after another, until the residue was a natural group; but 
this residue cannot be assumed to be the type of Conferva L.; that name is to be 
abandoned as a nomen confusum. The remnant in question has become two genera, 
Tribonema Derbes and Solier, 1858, and Bumilleria Borzi, 1895. They are unbranched 
filaments, common in freshwater pools. From typical green algae of similar appear- 
ance they are distinguished in the first place by the presence in each cell of several 
disk-shaped plastids without pyrenoids or with obscure ones. The cell walls, when 
treated with sulfuric acid, can be seen to consist of two parts like a barrel sawed 
across the middle. A broken filament ends always with a broken half wall. Monocilia, 
an unfamiliar alga isolated from soil, difi^ers in producing branching filaments. 

Phylum Phaeophyta [ 67 

Family 8. Phyllosiphonacea [Phyllosiphonaceae] Wille in Engler and Prantl. Nat. 
Pflanzenfam. I Teil, Abt. 2: 125 (1890). Family Vaucheriaceae (Nageli) Areschoug 
(1850), preoccupied by order Vaucheriaceae Nageli. Family Botrydiaceae Luther 
(1899). Heterokonta whose bodies are highly multinucleate single cells, filamentous 
or anchored by filamentous rhizoids. Botrydium is found on damp soil as dark green 
globes, sometimes as much as 2 mm. in diameter, anchored by much-branched color- 
less rhizoids. Vaucheria is a familiar alga on damp earth or in fresh water. It consists 
of irregularly branching filaments, green where exposed to light, colorless where 
growing downward and serving as rhizoids. The reproductive cells are cut off by 
walls. The end of an aerial filament, cut off in this fashion, may as a whole act as a 
spore. In water, the protoplast of such a cell may escape as an exceptionally large 
zoospore with as many pairs of flagella as the nuclei within it. Antheridia are brief 
branches, each releasing many minute sperms each with two unequal flagella. 
Oogonia are globular cells, multinucleate during development, but containing only 
one functional nucleus when mature. Phyllosiphon is of much the same structure as 
Vaucheria, but is parasitic in seed plants, particularly Araceae. It reproduces, ap- 
parently, only by the breaking up of the protoplast to produce minute non-flagellate 

Order 4. ChoanoflageUata [Choano-Flagellata] Kent Man. Inf. 1: 36 (1880). 
Order Bicoecidea Grasse and Deflandre in Grasse Traite Zool. 1, fasc. 1: 599 

Non-pigmented flagellates, usually attached, each cell bearing a single flagellum 
of the type called pantacroneme, with lateral appendages and a terminal whip-lash; 
the cell bearing also a protoplasmic collar, usually surrounding the base of the flagel- 
lum. The collar is a means of nutrition. Bacteria and other scraps of organic matter, 
driven against it by the beating of the flagellum, adhere and are carried to the interior 
of the cell by flow of the cytoplasm of which it consists. 

It is probable that the pantacroneme flagellum is a variant of the pantoneme 
flagellum, and that this order belongs naturally in class Heterokonta. It may have 
evolved from Silicoflagellata; or it may be that the collar is a modified flagellum, 
and that the group evolved from order Ochromonadalea. 

Most authors have recognized more than one family of choanoflagellates, but 
genera are not very numerous and one family seems sufficient to accommodate them. 

Family Bicoekida Stein Org. Inf. 3, I Halfte: x (1878). Family Craspedornona- 
dina Stein 1. c. Families Bikoecidae, Codonosigidae, Salpingoecidac, and Phalansteri- 
idae Kent op. cit. Families Codonoecina and Bikoecina Biitschli in Bronn Kl. u. Ord. 
Thierreichs 1: 814, 815 (1884). Families Bicoecaceae, Craspedoynonadaceae, and 
Phalanasteriaceae Senn in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. la: 121, 
123, 129 (1900). Family Gymnocraspedidae Grasse Traite Zool. 1, fasc. 1: 590 
(1952). Characters of the order. Cells naked, solitary: Monosiga; colonial: Codo- 
siga James-Clark [Codonosiga Stein), Sphaeroeca. Cells imbedded in gelatinous 
matter, the collars contracted: Phalanseterium. Loricate: Salpingoeca, Bicosoeca, 

The choanoflagellates were discovered by James-Clark (1866, 1868), who made at 
the same time the discovery that certain internal cavities of sponges are lined by 
minute cells (choanocytes) of the same structure as the choanoflagellates. From 
these observations he drew the conclusion that sponges are a sort of flagellates dis- 
tinguished by the production of exceptionally large and elaborate colonies. Kent 


The Classification of Lower Organisms 

described Proterospongia Haeckeli as a colonial organism of amoeboid and choano- 
flagellate cells in a common matrix; he regarded it as a transitional form, important 
as evidence of the evolution of sponges from choanoflagellates. According to Duboscq 
and Tuzet ( 1937) it is no organism, but a stage in the development of an individual 
sponge from one which has been damaged. In spite of this, the hypothesis that the 
choanoflagellates represent the evolutionary origin of the sponges, and accordingly 
of the entire animal kingdom, continues to appear tenable. 

Fig. 11. — Choanoflagellata : a, b, Monosiga spp.; c^ Phalanasterium digitatum; 
A, Salpingoeca ampullacea; e, Salpingoeca Clarkii; i, Foteriodendron petiolatum. 
c X 500, the remainder x 1,000. c-f after Stein (1878). 

Phylum Phaeophyta [ 69 

Order 5. Hyphochytrialea [Hyphochytriales] Bessey Morph. and Tax. Fungi 69 
Order Anisochytridiales Karling in American Jour. Bot. 30: 641 (1943), not 
based on a generic name. 

Non-pigmented organisms with walled cells, parasitic or saprophytic, the proto- 
plasm with numerous granules not of a shining appearance, producing zoospores 
with single anterior pantoneme flagella. 

The naked zoospores come to rest upon appropriate hosts or substrata. Ordinarily, 
in parasitic species, the protoplast of the zoospore makes its way to the interior of a 
cell of the host. It swells and develops a thin wall. The resulting structure may be 
called a center. In most members of the group, the center gives rise to a system of 
slender rhizoids; in some species, these give rise to further centers like the original 
one. Karling studied the cytology particularly in Anisolpidium. There are repeated 
simultaneous mitoses in the growing centers. Resting nuclei contain conspicuous 
karyosomes. Dividing ones show about five chromosomes in an intranuclear spindle 
which ends sharply in centrosomes. Eventually, in the usual course of events, each 
center produces an exit tube to the exterior. Its contents are released by delique- 
scence of the tip of the exit tube. Either before this or afterward, the mass of proto- 
plasm undergoes cleavage into uninucleate protoplasts which generate flagella. Some- 
times, instead of discharging their contents, the centers are converted into resting 
spores by the secretion of thick walls (this has been observed in only a few of the 
species). The resting spores germinate by producing exit tubes and discharging 
zoospores as ordinary centers do. 

The body type which has just been described may be called the chytrid body 
type; organisms of this body type were formerly assembled as a taxonomic group 
typified by the genus Chytridium. Couch, however, showed that these organisms 
form three groups distinguished by fundamental differences in type of flagellation. 
The present group is here given a place implying relationship to order Silicoflagellata. 

Karling (1943) accounted for fourteen species. He provided three families; only 
one is here maintained. 

Family Hyphochytriacea [Hyphochytriaceae] Fischer in Rabenhorst Kryptog.-Fl. 
Deutschland 1, Abt. 4:131 (1892). Families Anisolpidiaceae and Rhizidiomycetaceae 
Kailing in American Jour. Bot. 30: 641, 643 (1943). Characters of the order. 
Without rhizoids: Anisolpidium on brown algae; Roesia on Lemna; Cystochytrium 
on roots of Veronica. With rhizoids from a single center: Rhizidiomyces and Latr os- 
tium on green algae, aquatic fungi, and the empty exoskeletons of insects. With 
multiple centers: Hyphochytrium and Catenariopsis, on fungi and other hosts. 

Class 2. BACILLARIACEA Engler and PrantI 

Homalogonata Lyngbye Tent. Hydrog. Danicae 177 (1819). 
Order Diatomeae C. Agardh Syst. Alg. xii (1824). 

Division (of order ^/gae) Diatomaceae Harvey in Mackay Fl. Hibem. 166 (1836). 
Family Bacillaria Ehrenberg Infusionsthierchen 136 (1838). 
Series (of class Algae) Diatomaceae Harvey Man. British Alg. 15 (1841). 
Abtheilung (of cldiS,?, Isocarpeae) Diatomaceae Kiitzing Phyc. Germ. 54 (1845). 
Stamm Diatomea Haeckel Gen. Morph. 2: xxv (1866). 

Division (of class Algae) Diatomaceae Rabenhorst Kryptog.-Fl. Sachsen 1: 1 


The Classification of Lower Organisms 

Fig. 12. — Hyphochytrialea: a-e^ Anisolpidium Ectocarpii; a-c, individuals de- 
veloping in cells of Ectocarpus; d, mitotic figures x 2,000; e^ cell of Ectocarpus filled 
by a mature individual discharging spores, f, g, Rhizidiomyccs apuphysatus; f, zoo- 
spore; g, oogonium of Achlya parasitized by three individuals, h, i, j^ llyphochy- 
trium catenoides; h, zoospore; i, young individual; j, mature individual with fila- 
ments, sporangia, and zoospores in various stages of development. All after Karling 
(1943, 1944, 1939). x 1,000 except as noted. 

Phylum Phaeophyta [71 

Class Bacillariaceae Engler and Prantl Nat. Pflanzenfam. II Teil; 1 (1889). 

Subdivision and class Bacillariales Engler Syllab. 6 (1892). 

Hauptclasse Diatomeae Haeckel Syst. Phylog. 1: 90 (1894). 

Subclass Bacillariales Engler in Engler and Prantl Nat. Pflanzenfam. Teil I, Abt. 
la: V (1900). 

Class Bacillarieae Wettstein Handb. syst. Bot. 1: 74 (1901). 

Class Bacillarioideae Bessey in Univ. Nebraska Studies 7: 283 (1907). 

Class Diatomeae Schaffner in Ohio Naturalist 9: 447 (1909). 

Abteilung Bacillariophyta Engler. 

Ahteilung (of Stamm Chrysophyta) Diatomeae Pascher in Beih. bot. Centralbl. 
48, Abt. 2: 324 (1931). 

Class Bacillariophyceae Auctt. 

Unicellular (occasionally filamentous or colonial) organisms without flagella in 
the vegetative condition, each cell with one, two, or more plastids, brown, varying to 
yellow or exceptionally to bluish or colorless, and bearing a siliceous shell of two 
parts. Globules of oil and granules of something called volutin (the "red granules 
of Biitschli," apparently protein) are present. Other granules in some examples are 
said to be of leucosin. 

These organisms, the diatoms, are very common. There are some 5300 species. 
Microscopic examination of the bottoms of fresh water ponds reveals usually more of 
diatoms than of any other kind of organisms. Diatoms are frequent prey of many kinds 
of predators, from amoebas to whales. In using fish-liver oils as a source of vitamin 
D, man adds himself to a long chain of predators of which it is believed that diatoms 
are the usual ultimate prey. 

The shells of diatoms are not subject to decay. In certain places which were in 
the geologic past arms of the sea, there are enormous deposits of diatom shells in 
the form of a white earth. The oldest deposits are of the Cretaceous age. Thus it ap- 
pears that diatoms are a modern offshoot, no more ancient than the flowering plants. 
Diatomaceous earth is mined for various uses. It is an effective insulating material, 
and was the inert material first used in connection with nitroglycerine in the manu- 
facture of dynamite. 

The two parts of the shell of a diatom are called valves. They fit one over the 
other "like the parts of a pill box" (ZoBell, 1941, objects to this traditional simile, 
on the ground that in current language a pillbox is a concrete structure with loop- 
holes). The shells consist basically of something of the nature of pectin heavily im- 
pregnated with silica and characteristically sculptured. The cells appear markedly 
different in different aspects: the aspect which is in effect top or bottom view is 
called valve view, and that which is in effect side view is called girdle view. When a 
cell divides, each of the daughter cells receives one of the valves and generates an 
additional valve fitting within it. Diatoms in culture undergo a gradual diminution 
in size; there is an old hypothesis that this is caused by the fact that one of each pair 
of sister cells receives a slighly smaller valve than the other. 

Lauterborn (1896) described mitosis in Surirella and other diatoms. He found a 
centrosome, with radiating strands, near the nucleus. At the beginning of mitosis, 
the centrosome generates a disk-shaped structure which enters the nucleus and grows 
in such fashion as to become a cylinder extending through it. The cylinder is recog- 
nizably a spindle, but the chromosomes, instead of appearing within it, form a ring- 
shaped mass about its middle and divide into two ring-shaped masses which move 
along it to its extremities. The nuclear membrane ceases to be recognizable early in 

The Classification of Lower Organisms 

Fig. 13. — Bacillariacea : a, Mclosira sp., a living cell and an empty one. b, c. 
Girdle and valve views of cell of Cyclotella sp. d, e. Sections of a valve of Pinnu- 
laria sp., highly magnified, after Otto Miillcr (1896); d, about half-way between the 
middle and the end, e^ near the end. f, g, Girdle and valve views of Synedra sp. 
h, i. Girdle and valve views of Rhoicosphenia curvata. j, k, Girdle and valve views 
of Navicula sp. 1^ m, Girdle and valve views of Gomphonema sp. (the former show- 
ing the gelatinous stalk by which the cell is attached), n, o. Girdle and valve views 
of Cymbella sp. p, q, Surirella saxonica after Karsten (1900); p, two cells joined 
before conjugation; q, zygote; x 250. r, s, Girdle and valve views of Cocconeis sp. 
X 1,000 except as noted. 

Phylum Phaeophyta [ 73 

the process, but the nuclear cavity remains distinct until the chromosomes have 
reached the ends of the spindle. The nuclear sap and the spindle are then absorbed 
by the cytoplasm, but not until the spindle has budded off a new centrosome from 
each end. 

Subsequent authors, as Karsten (1900), Geitler (1927), Iyengar and Subrahman- 
yan (1942, 1944), and Subrahmanyan (1947), have not seen as full a series of stages 
as Lauterbom did. They have found centrosomes in at least some diatoms, and have 
confirmed the point that the spindle is a cylinder which is surrounded by the 
chromosomes instead of including them. 

The same authors have described sexual processes in Surirella, Cymbella, Coc- 
coneis, Cyclotella, and Navicula. In Surirella saxonica as described by Karsten, pairs 
of the wedge-shaped cells become attached by little bodies of slime at the narrow 
ends. Each nucleus divides twice, producing four, of which three are digested by the 
cytoplasm. The two protoplasts then move in amoeboid fashion out of their shells 
and they and their nuclei unite. The zygote protoplast grows to a size much greater 
than that of the parent cells and secretes a membrane which becomes silicified. The 
resulting cell is called an auxospore. 

In most kinds of diatoms, each cell produces two gametes. In some, the cells pair 
and proceed to produce auxospores individually, without conjugation. Karsten sup- 
sposed the latter examples to represent a stage in the evolution of sexual reproduc- 
tion under some zwingender Nothwendigkeit: much more probably, they are pro- 
ducts of degeneration. In Cyclotella, Iyengar and Subrahmanyan found the produc- 
tion of auxospores to involve autogamous karyogamy: the nucleus of a solitary cell 
undergoes meiosis; two of the haploid nuclei are digested, and the two which remain 
fuse with each other. It is evident that all diatoms are diploid in the vegetative 

The filamentous green Heterokonta Tribonema and Bumilleria are closely similar 
to the diatom Melosira, and it may reasonably be supposed that they represent the 
evolutionary origin of the group. 

Diatoms are preserved for study by violent methods which destroy the protoplasts, 
and the classification is based strictly on characters of the shells. So uniform is the 
group that Schiitt (in Engler and Prantl, 1896) treated it as a single family. He pro- 
vided an elaborate subsidiary classification involving two main groups. Subsequent 
scholars have found his system essentially sound as a representation of nature, but 
have raised the main groups to the rank of orders and the minor ones in correspond- 
ing degree. 

Order 1. Disciformia [Disciformes] Kiitzing Phyc. Germ. 112 (1845). 
Order Appendiculatae Kiitzing 1. c. 
Centricae Schiitt in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. lb: 57 

Order Centricae Campbell Univ. Textb. Bot. 90 (1902). 
Order Eupodiscales Bessey in Univ. Nebraska Studies 7: 284 (1907). 
Diatoms basically of radial symmetry, which, however, is often distorted; not 
motile in the vegetative condition; plastids numerous in the cells. 

These are the more primitive diatoms. The majority are marine. Three types of 
reproductive cells are known to be produced by them. 

Occasionally, in mass catches of material from the ocean, diatoms are found 
whose protoplasts have undergone repeated division within the shell and produced 

74 ] The Classification of Lower Organisms 

numerous little naked protoplasts. These protoplasts are said to bear flagella; whether 
one or two, equal or unequal, is not certainly known. They are supposed to escape 
and function as zoospores, but Karsten (1904), on rather scant evidence, supposed 
them to be gametes. 

A protoplast may contract and form a shell within its former shell. The new shell 
consists like the old one of two parts, one fitting within the other. The outer shell is 
usually more or less elaborately sculptured, while the inner is smooth. It is supposed 
that the outer shell is deposited between outer and inner masses of protoplasm, and 
that the entire protoplast then withdraws to the interior and deposits the inner shell 
in the opening. It is in this manner that the statospores of chrysomonads are formed. 
The resting cells of diatoms as just described are believed to be homologous with 
them, and are called by the same term. 

As a third manner of producing a reproductive cell, a protoplast may expand, force 
apart the valves of its shell, and deposit an enlarged shell about itself. The resulting 
spore is called an auxospore. As noted, Iyengar and Subrahmanyan found the pro- 
duction of auxospores in Cyclotclla to involve sexual processes. 

Schiitt divided the Centricae into three groups with names in -oideae (presum- 
ably subfamilies) and these into nine groups with names in -eae (presumably tribes). 
Subsequent authorities have made of Schiitt's groups a varying number of families. 
The minimum tenable number of families is three, corresponding to Schutt's 

Family 1. Coscinodiscea [Coscinodisceae] Kiitzing Phyc. Germ. 112 (1845). 
Family Melosireae Kiitzing op. cit. 66. Families Melosiraceae and Coscinodiscaceae 
West British Freshw. Alg. 274, 276 (1904). Melosira, in fresh water, the shells feebly 
silicified, the cells joined end to end in filaments. Cyclotclla, separate drum-shaped 
cells in fresh water. Coscinodiscus, the cells disk-shaped. Triceratium, cells of the 
form of 3-, 4-, or 5-sided prisms with abbreviated axes. 

Family 2. Rhizosoleniacea [Rhizosoleniaceae] West British Freshw. Alg. 278 
(1904). The cells, circular or elliptic in cross section, becoming elongate by inter- 
calation of ring-shaped bands of wall between the valves. Rhizosolenia. Corethron. 

Family 3. Biddulphiea [Biddulphieae] Kutzing Phyc. Germ. 115 (1845). Families 
Biddulphiaceae and Chaetoceraceae Auctt. Cells laterally compressed, elliptic in 
valve view, oblong or rhombic in girdle view. Cells of Biddulphia, solitary or colonial, 
are familar as epiphytes on marine algae. Chaetoceros, the cells with a long spine at 
each corner, frequently united valve to valve in filaments, abundant in subpolar 

Order 2. Diatomea [Diatomeae] C. Agardh Syst. Alg. xii (1824). 

Tribe Striatae with orders Astomaticae and Stomaticae, and tribe Vittatae also 

with orders Astomaticae and Stomaticae, Kutzing Phyc. Germ. ( 1845). 
Pennatae Schiitt in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. lb: 101 

Order Pennatae Campbell Univ. Textb. Bot. 90 ( 1902). 
Order Naviculales Bessey in Univ. Nebraska Studies 7: 284 ( 1907). 
Diatoms basically of isobilateral symmetry, occasionally so skewed as to be dorsi- 
ventral or asymmetric; valves usually punctured by a longitudinal cleft called the 
raphe, or bearing a marking of some sort, called the pseudoraphe, in the same posi- 
tion; exhibiting, when possessed of a true raphe, a gliding motion; cells usually with 
two plastids. 

Phylum Phaeophyta [ 75 

The motion of the pennate diatoms is a gliding upon surfaces, with frequent re- 
versal, in either direction of the long axis of the cell. It depends upon the flow of a 
stream of exposed protoplasm. This is the opinion of Max Schultze (1865), Otto 
Miiller (1889, 1896), and Lauterborn (1896); there have been other hypotheses. 
Miiller showed that the true raphe, without which the motion does not occur, is an 
actual opening. The raphe is not a simple crack; it enters the wall obliquely and 
bends at a sharp angle to come from another oblique direction to the interior. Its 
proportions vary along its length, and it is interrupted at the middle of the valve by 
a knob, the central granule, projecting inward from the valve. 

The pennate diatoms do not produce flagellate cells nor statospores, but they pro- 
duce auxospores, usually by sexual processes. The majority inhabit fresh water. 

Eleven families are currently recognized. 

a. Without raphes. 

Family 1. Fragilariea [Fragilarieae] (Harvey) Kutzing Phyc. Germ. 62 (1845). 
Family Fragilariaceae West British Freshw. Alg. 285 (1904). Cells symmetrical with 
respect to three planes, without internal partitions. Fragilaria. Synedra. 

Family 2. Tabellariea [Tabellarieae] Kutzing op. cit. 110. Family Tahellariaceae 
West op. cit. 281. Cells symmetrical with respect to three planes, with longitudinal 
internal partitions. Tabellaria. 

Family 3. Bacillaria Ehrenberg Infusionsthierchen 136 (1838). Family Diato- 
maceae West op. cit. 284. Cells symmetrical with regard to three planes, with trans- 
verse internal partitions, solitary, or joined valve to valve in ribbons, or corner to 
comer in zig-zag chains. Diatoma. 

Family 4. Meridiea [Meridieae] Kutzing op. cit. 61. Family Meridionaceae West 
op. cit. 283. Cells symmetrical with regard to two planes, wedge-shaped both in valve 
and in girdle view, with transverse internal partitions, often joined valve to valve 
in fan-shaped colonies which are sometimes so extended as to produce spiral fila- 
ments. Meridion. 

b. With raphes, the valves of each cell alike. 

Family 5. Naviculea [Naviculeae] Kiitzing op. cit. 90. Family Naviculaceae Rab- 
enhorst Kryptog.-Fl. Sachsen 1: 33 (1863). This is the most numerous family of 
diatoms. In most of the genera the cells are narrowly rectangular in girdle view, 
narrowly elliptic in valve view, being of the shape of flat-bottomed boats. Navicula, 
Pinnularia, etc. In other genera, as Gyrosigma and Pleurosigma, the cells are so 
skewed as to be sigmoid in valve view. 

Family 6. Gomphonemea [Gomphonemeae] Kiitzing op. cit. 87. Family Gom- 
phonemaceae West op. cit. 297. Cells wedge-shaped. Gomphonema. 

Family 7. Cymbellea [Cymbelleae] (Harvey) Kiitzing op. cit. 84. Family Cocco- 
nemaceae West op. cit. 298. Cells with two planes of symmetry, in valve view crescent- 
shaped or approximately so. Cymbella. Rhopalodia. 

Family 8. Eunotiea [Eunotieae] Kiitzing op. cit. 57. Family Eunotiaceae West op. 
cit. 287. Cells curved as in the preceding family, the raphes reduced to brief clefts 
near the ends of the valves. Eunotia. 

Family 9. Nitzschiacea [Nitzschiaceae] West op. cit. 301. Cells asymmetric in 
valve view, the raphe along one margin. Nitzschia. Hantschia. 

Family 10. Surirellea [Surirelleae] Kiitzing op. cit. 70. Family Surirellaceae West 
op. cit. 303. Each cell with two marginal raphes. Surirella. 

c. The two valves of each cell unlike, one with a raphe, one with a pseudoraphe. 

76 ] The Classification of Lower Organisms 

Family 11. Achnanthea [Achnantheae] Kiitzing op. cit. 81. Families Achnan- 
thaceae and Cocconeidaceae West op. cit. 289, 290. Achnanthes, Rhoicosphenia, 

Class 3. OOMYCETES Winter 

Class OoMYCETEs Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 1: 32 

Phycomyceten de Bary Vergl. Morph. Pilze 142 ( 1884), in part. 
Class Phycojnycetes Engler and Prantl Nat. Pflanzenfam. II Teil: 1 (1889), in 

Reihe Oomycetes Fischer in Rabenhorst Kryptog.-FI. Deutschland 1, Abt. 4: 310 

Stamm Phykomycophyta Pascher in Beih. bot. Centralbl. 48, Abt. 2: 330 (1931), 

in part. 
Biflagellatae Sparrow Aquatic Phycomycetes 487 (1943). 

Organisms of fungal or chytrid body type, that is, non-pigmented saprophytes or 
parasites whose bodies are walled filaments or cells with or without rhizoids; the 
walls consisting partially of cellulose; reproducing asexually by zoospores with 
paired unlike flagella which are, so far as is known, respectively pantoneme and 
acroneme, and usually sexually by fertilization, the eggs being distinct cells within 
the oogonia. The regularly cited example and evident standard genus of the group 
is Saprolegnia. 

Conventional botanical classification recognizes within the group of Fungi a sub- 
ordinate group named Phycomycetes, which is in turn divided into Oomycetes and 
Zygomycetes, the former including the chytrids. This arrangement suggests an evo- 
lutionary series, originating perhaps among non-pigmented flagellates, and leading 
through chytrids, typical Oomycetes, and Zygomycetes to the typical fungi. It does 
not now appear tenable. Couch (1939) pointed out differences between Oomycetes 
and Zygomycetes which make any direct connection between them appear quite 
improbable; and his observations on flagella showed that only a small minority among 
organisms of chytrid body type have anything to do with the proper Oomycetes. 

There is an old hypothesis (Sachs, 1874) that Vauchcria may represent the direct 
ancestry of Saprolegnia. This hypothesis could not be taken seriously while Sapro- 
legnia and its allies were known to produce heterokont zoospores, while Vaucheria 
was supposed to be a typical isokont green alga. Now it again appears probable. It 
implies that in the present group the fungal body type is more primitive than the 

The Oomycetes may be organized as three orders. 
l.Of fungal body type, i.e., consisting of fila- 

2. Essentially aquatic Order 1. Saprolegnina. 

2. Mostly not aquatic, parasitic on higher 

plants Order 2. Peronosporina. 

1. Of chytrid body type, i.e., the cells not elong- 
ated to filamentous form, though sometimes 
proliferating or producing rhizoids Order 3. LAGENroiALEA. 

Phylum Phaeophyta [ 77 

Order 1. Saprolegnina [Saprolengninae] Fischer in Rabenhorst Kryptog.-Fl. 
Deutschlandl,Abt.4: 311 (1892). 
Order Eremospermeae and suborder Mycophyceae Kiitzing Phyc. Gen. 146 

(1843), in part. 
Order Oosporeae Cohn in Hedwigia 11: 18 (1872), in part. 
Order Oomycetes and suborder Saprolegniineae Engler Syllab. 24 (1892). 
Order Saprolegniineae Campbell Univ. Textb. Bot. 153 (1902). 
Order Siphonomycetae Bessey in Univ. Nebraska Studies 7: 286 (1907). 
Order Saprolegniales Auctt. 

Order L^p^omzfa/^?^- Kanouse in American Jour. Bot. 14: 295 (1927). 
Aquatic Oomycetes, filamentous, saprophytic or facultatively parasitic, the zoo- 
spores diplanetic (exhibiting two periods of swimming) or giving evidence of an 
ancestral diplanetic condition. The old ordinal names Eremospermeae and Oosporeae 
designated miscellaneous collections of groups in which this one was listed at or near 
the beginning. Either one, if taken up, would be applied here, but it seems better to 
treat them as nomina confusa. 

1. Filaments not constricted Family 1. Saprolegniea. 

1. Filaments constricted at intervals. 

2. Filaments not differentiated into basal 

and reproductive parts Family 2. Leptomitea. 

2. Filaments differentiated into basal and 

reproductive parts Family 3. Rhipidiacea. 

Family 1. Saprolegniea [Saprolegnieae] Kiitzing Phyc. Gen. 157 (1843). Family 
Saprolegniaceae Cohn in Hedwigia 11 : 18 (1872). Aquatic Oomycetes consisting of 
branching filaments of essentially uniform diameter without crosswalls other than 
those which set apart differentiated reproductive structures. 

These well-known organisms are called water molds. According to Coker (1923) 
there are about eighty definitely recognizable species. They may be parasitic on 
fishes or saprophytic on organic remains in water or soil. In almost any body of soil 
or of fresh water they may be found by "baiting," in former practice with dead flies, 
currently with hemp seeds. 

Mitosis has rarely "been observed in the vegetative filaments, the nuclei being very 
minute. Eggs are produced in large globular multinucleate oogonia borne at the ends 
of filaments. The nuclei in the developing oogonia become enlarged and undergo a 
single flare of concurrent mitoses (Davis, 1903; Couch, 1932). The sharp-pointed 
spindles, ending in centrosomes, are formed within the nuclear membrane. The 
membrane disappears toward the end of the mitotic process, and a nucleolus, which 
has persisted to this stage, undergoes solution in the cytoplasm. The chromosome 
numbers (Ziegler, 1953) are 3, 4, 5, 6, or 7. 

Within each oogonium there appear one or a few minute bodies called coenocentra. 
One nucleus becomes associated with each coenocentrum; all others break down and 
disappear. Each surviving nucleus with the cytoplasm associated with it becomes 
organized as an egg. When several eggs are produced, they share all of the cytoplasm 
of the oogonium; when only one egg is produced, some of the cytoplasm is left out- 
side of it. 

Sperms are produced in small multinucleate antheridia borne at the tips of fila- 
ments in contact with oogonia. Typically, each individual bears both oogonia and 
antheridia. Some species are capable of self-fertilization; others exist as two kinds 
of individuals, each capable of fertilizing the other; some occur as distinct male and 


The Classification of Lower Organisms 


' •ITi''!' nVgi 'Villi 

Fig. 14. — Oomycetes: a. Filaments and sporangia of Dictyuchus sp. x 50. 
b, C, Zoospores of the second stage of swimming, of Achlya caroliniana and Sapro- 
legnia ferax, after Couch (1941) x 1,000. d^ Oogonia and antheridia of Dictyuchus 
X 400. e, f, g, Saprolegnia mixta after Davis (1903) : e, developing oogonium with 
numerous nuclei x 500; f, metaphase of nuclear division x 2,000; g, developing 
oogonium in which most of the nuclei have undergone degeneration; a few have 
become associated with coenocentra, and the cytoplasm is undergoing cleavage to 
produce eggs about these. 

Phylum Phaeophyta [ 79 

female individuals. Parthenogenesis (reproduction by eggs which have not been 
fertilized) is rather common in this group. There are no swimming sperms: nuclei 
from the antheridia reach the eggs through fertilization tubes, or by migration through 
the periplasm. 

Ziegler found that the first nuclear divisions of the nucleus of the zygote are 
meiotic: all cells except the zygotes are haploid. 

The organs of asexual reproduction are cylindrical sporangia terminal on the fila- 
ments. Within these the multinucleate protoplasts undergo cleavage into minute 
uninucleate spores. It is chiefly by details of the behavior of the sporangia and spores 
(the latter diplanetic, monoplanetic, or not swimming at all) that the dozen genera 
are distinguished. Diplanetism is the character of zoospores which are not directly 
infective; they undergo encystment, and the cysts release infective zoospores. During 
the first stage of swimming, the spores are pear-shaped, with the nucleus drawn out 
into a beak toward the narrow anterior end, where the flagella are attached. Spores re- 
leased from cysts for a second period of swimming are bean-shaped, with the flagella 
attached laterally, each connected through a separate rhizoplast to the nucleus, which 
lies at some distance from the cell membrane (Cotner, 1930). No explanation of 
this behavior, whether by phylogeny, genetics, physiology, or competitive advantage, 
is known. The apparent trend of evolution is to eliminate it. Monoplanetic spores 
in the present group are usually released from the sporangia as naked protoplasts 
which undergo encystment and emerge subsequently as flagellate spores of the second 

Saprolegnia releases diplanetic spores through circular pores in the tips of sporangia 
in which the spores are formed in several rows; new sporangia develop within empty 
old ones. Organisms which differ from Saprolegnia only in producing new sporangia 
beside, instead of within, the old ones, were formerly assigned to Achlya, but are now 
called Isoachlya. Leptolegnia differs from Saprolegnia and Isoachlya in forming 
spores in a single row. In Achlya proper, the spores are discharged without flagella, 
to encyst and swim only once. In Thraustotheca the monoplanetic spores are re- 
leased by irregular breakdown of the distal part of the sporangium. In Dictyuchus 
the spores become encysted before discharge; their protoplasts escape in the form of 
secondary swarmers through individual pores in the wall of the sporangium. Salvin 
(1942) found that cultures while growing release into the medium substances which 
affect the type of sporangium produced, so that a given culture may be while young 
of the character of Achlya, and later of the character of Thraustotheca or Dictyuchus. 

Family 2. Leptomitea [Leptomiteae] Kiitzing Phyc. Gen. 150 (1843). Family 
Leptomitaceae Schroter in Engler and Prantl Nat. Pflanzenfam. I Tail, Abt. 1 : 101 
(1893). Oomycetes consisting of filaments which are constricted at intervals, but 
are not differentiated into a basal cell and reproductive branches. In sewage or on 
organic matter decaying in water. Leptomitus, Apodachlya, Apodachlyella, with 
some seven known species. The numbers of species and degree of distinction of this 
family and the following do not appear to justify the proposed establishment of a 
separate order for them. 

Family 3. Rhipidiacea [Rhipidiaceae] Sparrow in Mycologia 34: 116 (1942). 
Saprophytes resembling the Leptomitea, the body differentiated into a main part, the 
basal cell, rhizoids of limited growth, and slender branches bearing the reproductive 
structures. Sapromyces, Araiospora, Rhipidium, Mindeniella, with perhaps a dozen 
known species. 

80 ] The Classification of Lower Organisms 

Order 2. Peronosporina [Peronosporinae] Fischer in Rabenhorst Kryptog.-Fl. 
Deutschlandl,Abt.4: 383 (1892). 
Suborder Peronosporineae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, 

Abt. 1: iv (1897). 
Order Peronosporineae Campbell Univ. Textb. Bot. 155 (1902). 
Order Peronosporales Auctt. 
Mostly parasites on terrestrial plants, but including also aquatic parasites and a 
few saprophytes, the bodies filamentous, reproducing sexually by fertilization, the 
eggs solitary in the oogonia, reproducing asexually chiefly by conidia, that is, by air- 
born cells cut off from the ends of the filaments. The conidia are homologous with 
the sporangia of the Saprolegnina : they germinate in most examples by release of 
zoospores (which show no signs of diplanetism), but in the more highly evolved 
examples they give rise to filaments. Ferris (1954) found the zoospores of Phytoph- 
thora to bear the paired flagella, respectively pantoneme and acroneme, which are 
typical of Phaeophyta. 

In the multinucleate oogonia of most members of the group, single flares of mitoses 
occur. The sharp-pointed spindles, described in some accounts as ending in centro- 
somes, are formed within the persistent nuclear membrane, which undergoes con- 
striction during the final stages of mitosis. A coenocentrum appears (this structure 
was first described as occurring in Albugo, by Stevens, 1899); in general, one nucleus 
becomes associated with it, and is thus selected as the egg nucleus, the remaining 
nuclei being cast out to undergo disolution in a body of periplasm. The antheridium 
develops in contact with the oogonium, and fertilization is accomplished by the 
growth of a fertilization tube through the periplasm to the egg (Davis, 1900; Stevens, 

In Albugo Bliti and A. Tragopogonis, Stevens observed two flares of simultaneous 
mitoses in the oogonium and antheridium. If this phenomenon were general in the 
group one would confidently identify it as meiosis. The single coenocentrum attracts 
many nuclei; the fertilization tube delivers a large number of sperm nuclei; thus 
multiple karyogamy occurs within a single cell. The further history of the resulting 
peculiar zygote, containing many nuclei which are not by any evident necessity 
genetically uniform, is unknown. 

This order is evidently a specialized offshoot of the preceding. The family Pythiacea 
is a good example of a transition group; many authorities have assigned it to the pre- 
ceding order. 

1. Producing solitary globular sporangia or 
conidia at the ends of scarcely specialized 

filaments; mostly aquatic Family 1. Pythiacea. 

1. Producing conidia usually in clusters at the 
ends of specialized filaments (conidio- 
phores) ; parasites on land plants. 

2. Conidiophores brief, unbranched, the 

conidia in chains Family 2. Albuginacea. 

2. Conidiophores elongate, usually branch- 
ed, the conidia solitary or clustered, not 

in chains Family 3. Peronosporacea. 

Family 1. Pythiacea [Pythiaccae] Schroter in Engler and Prantl Nat. Pflanzenfam. 
I Teil, Abt. 1: 104 (1893). Aquatic parasites and saprophytes releasing zoospores 
from globular reproductive structures terminal on the filaments, together with para- 

Phylum Phaeophyta [81 

sites attacking land plants under moist conditions. The reproductive structures act 
as sporangia if formed in water, as conidia if formed in air. Pythium, saprophytic on 
plant remains in water or parasitic on algae or higher plants, includes some forty 
species (Matthews, 1931). The few other genera include perhaps a dozen species. 
Zoophagus produces specialized branches which serve as traps for rotifers which are 
parasitized and killed. 

Family 2. Albuginacea [Albuginaceae] Schroter op. cit. 110. Parasites of higher 
plants, called white rusts, the masses of conidia which push up and burst through the 
epidetmis being of a white color. Albugo. 

Family 3. Peronosporacea [Peronosporaceae] Cohn in Hedwigia 11: 18 (1872). 
Parasites of higher plants, called downy mildews. The ovoid conidia are produced 
solitary or in clusters, not in chains, on elongate conidiophores, usually branched, 
projecting through the stomata of the hosts. This numerous group includes the 
agents of some of the most important diseases of cultivated plants. Plasmopara viti- 
cola, causing downy mildew of grapes. Phytophthora injestans, the cause of the blight 
of potatoes which produced the Irish famine of 1846. Peronospora, the many species 
attacking many kinds of plants. 

Order 3. Lagenidialea [Lagenidiales] Karling in American Jour. Bot. 26: 518 
Suborder Ancylistineae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, 

Abt. 1 : iv ( 1897), for the most part, not as to the type genus Ancylistes. 
Order Ancylistales Auctt., in part. 
Oomycetes of chytrid body type, parasites consisting of walled cells which are 
more or less isodiametric, sometimes proliferating or producing rhizoids, but not 
forming extensive branched filaments. The cells become multinucleate. Mitotic 
figures of Olpidiopsis as described by Barrett (1912) and McLarty (1941) are quite 
as in the preceding orders, with sharp-pointed intranuclear spindles apparently with 
centrosomes at the poles. In the usual course of events, each cell develops an exit 
tube to the exterior of the host, and the protoplast becomes divided into uninucleate 
cells which escape as unequally biflagellate zoospores. Fertilization, by the migration 
of the protoplast of one cell into another, has been observed; the zygote becomes a 
thick-walled resting spore. 

1. Internal parasites without rhizoids. 
2. The cells not proliferating. 

3. The zoospores diplanetic Family 1. Ectrogellacea. 

3. The zoospores not diplanetic Family 2. Olpidiopsidacea. 

2. The cells proliferating. 

3. Marine Family 3. Sirolpidiacea. 

3. Fresh-water Family 4. Lagenidiacea. 

1. External parasites with rhizoids Family 5. Thrau stock ytriacea. 

Family 1. Ectrogellacea [Ectrogellaceae] Scherffel in Arch. Prot. 52: 6 (1925). 
Ectrogella, Eurychasma, Eurychasmidium, Aphanomycopsis, with about a dozen 
known species, attacking diatoms and red and brown algae. 

Family 2. Olpidiopsidacea [Olpidiopsidaceae] Sparrow in Mycologia 34: 116 
(1942). Olpidiopsis and a few other genera, with some thirty known species, attack- 
ing water molds, green algae, red algae, and other aquatic organisms. 

Family 3. Sirolpidiacea [Sirolpidiaceae] Sparrow 1. c. Sirolpidium and Pontisma, 
each with one species, attacking marine algae, respectively green and red. 

82 ] The Classification of Lower Organisms 

Family 4. Lagenidiacea [Lagenidiaceae] Schroter in Engler and Prantl Nat. 
Pfianzenfam. I Teil, Abt. 1 : 89 (1893). Lagenidium, Myzocytium, and Lagenocystis^, 
with some twenty known species, attacking green algae, rotifers, pollen which has 
fallen into water, and the roots of grasses. 

Family 5. Thraustochytriacea [Thraustochytriaceae] Sparrow op. cit. 115. The 
single species Thraustochytrium proliferum Sparrow was found as solitary cells ex- 
ternal on certain marine green algae and red algae which are penetrated by means 
of branching rhizoids. Reproduction is by release of naked protoplasts which become 
laterally biflagellate after a period of rest. 

Class 4. MELANOPHYCEA (Ruprecht) Rabenhorst 

Order Fucacees Lamouroux in Ann. Mus. Hist. Nat. Paris 20: 28 (1813). 

FucoiDEAE C. Agardh Synops. Alg. Scand. ix (1817). 

Order Fucoideae C. Agardh Syst. Alg. xxxv (1824). 

Division (of order Algae) Melanospermeae Harvey in Mackay Fl. Hibern. 157 

Series (of order Algae) Melanospermeae Harvey Man. British Alg. 1 (1841). 
Order Pycnospermeae and tribe Angiospermeae Kiitzing Phyc. Gen. 333, 349 

Class Fucoideae J. Agardh Sp. Alg. 1 : 1 (1848). 

Melanophyceae Ruprecht in Middendorff Sibir. Reise 1, part 2: 200 (1851). 
Class Melanophyceae Rabenhorst Kryptog.-Fl. Sachsen 1: 275 (1863). 
Stamm Fucoideae Haeckel Gen. Morph. 2: xxxv (1866). 
Series {Reihe) Phaeophyceae Hauck in Rabenhorst Kryptog.-Fl. Deutschland 2: 

282 (1885). 
Class Phaeophyceae Engler and Prantl Nat. Pfianzenfam. H Teil: 1 (1889), 
Class Dictyotales Engler in Engler and Prantl Nat. Pfianzenfam. I Teil, Abt. 2 : 

Classes Phaeosporeae, Tetrasporeae, and Cyclosporeae Bessey in Univ. Nebraska 

Studies 7: 288, 290 (1907). 
CXdiSS Dictyoteae Schaffner in Ohio Naturalist 9: 448 (1909). 
Subclass Melanophyceae Setchell and Gardner in Univ. California Publ. Bot. 8: 

387 (1925). 
Classes Isogeneratae, Heterogeneratae (with subclasses Haplostichinae and Poly- 
stichinae) and Cyclosporeae Kylin in Kungl. Fysiog. Sallsk. Handl. n. f. 44, no. 
7: 91 (1933). 
Filamentous or thallosc Phaeophyta, yellow to brown in color and living by photo- 
synthesis, producing reproductive cells with paired unequal flagella. 

These are the typical brown algae. They are almost exclusively marine, being 
abundant along with red and green algae on most coasts, and particularly abundant 
farther toward the poles than the red and green groups. The lower brown algae are 
branched filaments of microscopic dimensions, commonly epiphytic on other algae. 
More highly developed examples are thallosc and anchored to rocks. Some of these, 
particularly the ones whose English name is kelp, reach great sizes and considerable 
elaboration of structure. Papenfuss (in Smith, 1951) gives the number of genera as 
about 240, and that of known species as about fifteen hundred. 

^Lagenocystis nom. nov. Lagena Vanterpool and Ledingham in Canadian 
Jour Res. 2: 192 (1930), non Parker and Jones 1859. L. radicicola (Vanter- 
pool and Ledingham) comb. nov. 

Phylum Phaeophyta [ 83 

The cells are walled chiefly with readily hydrolyzable modified polysaccharides. 
Algin, the soda extract of kelps, consists of chains of oxidized mannose units. A poly- 
saccharide of the sugar fucose, with a sulfate radicle to each sugar unit, is also present. 
A small percentage of cellulose is present, apparently as the immediate investment 
of each protoplast. A glycogen- or dextrin-like dextrosan, laminarin, is stored (Miwa, 
1940; Tseng, 1945). The plastids contain chlorophylls a and c (Strain, in Franck 
and Loomis, 1949) and carotin; xanthophyll is also present in the more primitive 
examples. In all examples, there is an additional carotinoid called fucoxanthin, which 
produces the brown color. The analytic process of separating the pigments yields 
also a sterol, fucosterol, not found in green plants; but this substance, and fucoxanthin, 
are found in chrysomonads, green Heterokonta, and diatoms (Carter, Heilbron, and 
Lythgoe, 1939). 

Cytological study of a considerable variety of brown algae (Swingle, 1897; Farmer 
and Williams, 1896; Mottier, 1898, 1900; Simons, 1906; Yamanouchi, 1909, 1912; 
McKay, 1933) has shown that the spindle and chromosomes appear within an intact 
nuclear membrane which disappears during the later stages of division. A centrosome, 
usually with radiating rays, is present outside of the membrane at each pole of the 
spindle. In Stypocaulon, a comparatively primitive brown alga, Swingle found the 
centrosome to be a permanent structure, dividing as a preliminary to each division 
of the nucleus. In the generality of brown algae, the centrosomes appear de novo as 
division begins. 

Swimming cells are produced by primitive brown algae as spores and as morpholo- 
gically undifferentiated gametes; in the most advanced brown algae, such cells are 
produced only as sperms. The flagella are attached laterally. The anterior flagellum 
is the longer except in order Fucoidea (Kylin, 1916). Longest (1946) found in 
Ectocarpus that the anterior flagellum is pantoneme, and the posterior one acroneme. 
The swimming cells are without walls, and contain, beside the nucleus, usually one 
plastid and a light-sentitive speck, the stigma or eyespot. They are quite small. No 
system of structures linking the nuclei, centrosomes, and flagella has been discovered. 

Thuret (1850) discovered that most brown algae produce swimming cells from 
structures of two different sorts, which he named (1855) respectively plurilocular 
sporangia and unilocular sporangia. The difference between them is this. In the 
developing plurilocular structure, each division of the nucleus is followed by division 
of the protoplast and deposition of a wall, with the result that the swimming cells 
emerge from separate walled spaces. In the unilocular structure, the nucleus divides 
repeatedly before the protoplast divides; the protoplast then undergoes cleavage to 
produce swimming cells which emerge from a single walled space. A number of 
studies (Clint. 1927; Higgins, 1931; Knight, 1923, 1929) have shown that the first 
two nuclear divisions in the unilocular structure are normally meiotic. Unilocular 
structures occur normally only on diploid individuals and release haploid swimming 
cells. A few exceptional species, however, are known to bear unilocular structures 
which produce swimming cells without the intervention of meiosis. 

In Ectocarpus siliculosus as studied by Berthold (1881) at Naples, the swimming 
cells from unilocular structures are spores which give rise to haploid individuals. In 
the same species as studied in the Irish Sea by Knight (1929), they were found to 
act as gametes, conjugating and giving rise to diploid individuals. Diploid and hap- 
loid individuals of Ectocarpus are alike, and E. siliculosus may be said to have a 
facultatively complete homologous life cycle. The haploid individuals produce pluri- 
locular reproductive structures; the swarmers from these act either as spores, re- 


The Classification of Lower Organisms 

Fig. 15. — Stages of nuclear division in Stypocaulon x 1,000 after Swingle (1897). 

Phylum Phaeophyta [ 85 

producing the haploid stage, or as gametes, initiating the diploid stage. The diploid 
individuals produce both plurilocular and unilocular reproductive structures. The 
swarmers from the former are spores, reproducing the diploid body. The swarmers 
from the latter act either as spores, giving rise to haploid individuals, or as gametes, 
reproducing the diploid body. 

It is believed that the brown algae arose by evolution from order Ochromonadalea. 
Filamentous organisms with a facultatively complete homolgous life cycle, as just 
described, are believed to be primitive among them : such organisms appear to be the 
starting point of evolution in many features. The filaments have become differentiated 
and woven into thalli, and thalli of tridimensionally placed cells have been produced. 
The haploid and diploid stages have become differentiated. The plurilocular and 
unilocular structures have undergone specialization. Even in the most primitive 
brown algae, there is a physiological differentiation of gametes; this has evolved into 
extreme morphological differentiation. Every one of these evolutionary changes ap- 
pears to have occurred in more than one line of descent; research is constantly reveal- 
ing intermediate examples and rather free parallel evolution. 

Conservative classification, such as that of Fritsch (1945), recognizes as orders a 
comparatively primitive miscellany followed by a series of small derived groups 
marked by distinctive specializations. Features of the life cycle, as applied to classi- 
fication by Taylor (1922), Oltmanns (1922), Svedelius (1929) and Kylin (1933), 
are not reliable as marks of natural groups. Kylin provided three classes (one of 
them divided into two subclasses) and twelve orders. His system appears to provide 
an excessive number of subdivisions of high category within a moderately small group 
exhibiting no very profound evolutionary gaps. Tentatively, the seven orders dis- 
tinguished as follows may be recognized. 

1. Producing spores, that is, cells which germi- 
nate without syngamy. 

2. All spores bearing flagella. 

3. Having an alternation of haploid 
and diploid stages which are alike, 
both being filamentous; or else com- 
pletely lacking one of these stages. 

4. The filaments uniseriate Order 1. Phaeozoosporea. 

4. The filaments becoming pluri- 

seriate Order 2. Sphacelarialea. 

3. Not as above. 

4. Haploid stage thallose, not dis- 
tinctly less highly developed 

than the diploid stage Order 5. Cutlerialea. 

4. Haploid stage filamentous, dis- 
tinctly less highly developed 
than the diploid stage. 

5. Diploid stage filamentous; 
or, if partially or com- 
pletely thallose, the thal- 
lose part with apical growth Order 4. SpoROCHNoroEA. 

5. Diploid stage thallose, its 

growth intercalary Order 6. Laminariea. 

2. Producing large non-motile spores Order 3. Dictyotea. 

86 ] The Classification of Lower Organisms 

1. Producing no spores; all individuals diploid 

and reproducing exclusively sexually Order 7. FucoroEA. 

Order 1. Phaeozoosporea [Phaeozoosporeae] Hauck in Rabenhorst Kryptog.-Fl. 
Deutschland 2: 312 (1885). 
Order Syntamiidae Areschoug in Act. Reg. Soc. Upsala 14: 387 ( 1850) , in part; 

a nomen confusum. 
Order Ectocarpeae J. Agardh Sp. Alg. 1: 6 (1848), preoccupied by family 

EcTOCARPEAE KUtzing (1843). 
Section (of Algae Zoosporeac) Phaeosporeae Thuret in Ann. Sci. Nat. Bot. 

ser. 3, 14: 233 (1850). 
Order Phaeosporeae Wettstein Handb. syst. Bot. 1: 173 (1901). 
Order Ectocarpales Bessey in Univ. Nebraska Studies 7: 288 (1907). 
Order Phaeosporales and suborder Ectocarpineae Taylor in Bot Gaz. 74: 435, 
436 (1922). 
Microscopic brown algae of the form of undifferentiated uniseriate branching fila- 
ments, mostly with distinct haploid and diploid stages (exceptionally lacking the 
former), the stages distinguishable only by the limitation of unilocular reproductive 
structures to the diploid stage, the gametes morphologically uniform. 

The order is typified by Ectocarpus, which is by coincidence also the theoretical 
ancestral type of the brown algae, the living organism which supposedly represents 
the evolutionary origin of the group. Recent systems of classification limit this order, 
formerly construed as extensive, to this genus and a few others, as Pylaiella and Streb- 
lonema, which make up the family Ectocarpea [Ectocarpeae] Kiitzing (family Ecto- 
carpaceae Cohn). 

Order 2. Sphacelarialea [Sphacelariales] (Oltmanns) Engler and Gilg Syllab. ed. 
9 u. 10: 27 (1924). 
Order Sphacelarieae J. Agardh Sp. Alg. 1: 27 (1848), preoccupied by family 

Sphagelarieae Kiitzing (1843). 
Sphacelariales Oltmanns Morph. u. Biol. Alg. ed. 2, 2: 2 (1922). 
Brown algae distinguished from the Ectocarpea only by features of the vegetative 
structure, namely that the filaments have large apical cells, and that the cells cut off 
from them divide lengthwise without increasing considerably in thickness, with the 
result that the filaments consist of tiers of cells. The life cycle is the same as in Ecto- 
carpea. Family Sphacelariea [Sphacelarieae] Kiitzing (family Sphacelariaceae Cohn) 
includes Sphacelaria and Stypocaulon. A few other families have been segregated. 

Order 3. Dictyotea [Dictyoteae] Greville Alg. Brit. 46 (1830). 
Tribe Dictyoteae Harvey in Mackay Fl. Hibern. 159 (1836). 
Family Dictyoteae Kiitzing Phyc. Gen. 337 (1843). 
Order Dictyotaceae Hauck in Rabenhorst Kryptog.-Fl. Deutschland 2: 302 

Class Dictyotales Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 2 : 

ix (1897). 
Akinetosporeae Oltmanns Morph u. Biol. Alg. 1: 473 (1904). 
Order Tilopteridales and Class Tetrasporeae with order Dictyotales Bessey in 

Univ. Nebraska Studies 7: 290 (1907). 
Scries Aplanosporeae Setchell and Gardner in Univ. California Publ. Bot. 8: 

649 (1925). 

Phylum Phaeophyta [ 87 

Filamentous or thallose brown algae with haploid and diploid stages equally de- 
veloped, producing large spores without flagella, solitary or few in the sporangia. 

Here are placed two families, Tilopteridea and Dictyotacea. 

Family Tilopteridea [Tilopterideae] Cohn is a small group, apparently known 
only from European coasts. They are evidently closely related to the Ectocarpea. 
They consist of branching filaments which may become pluriseriate. In Haplospora 
(poorly known; but Tilopteris and other genera are even more so), the haploid stage 
bears both plurilocular structures, releasing minute swimming cells of the structure 
usual in brown algae, and unilocular structures which release their contents as single 
uninucleate protoplasts without flagella. The diploid stage bears only unilocular 
structures which release their contents as single quadrinucleate non-motile spores. It 
is inferred that the swimming cells from the plurilocular structures are sperms, and 
that the protoplasts released from the unilocular structures on haploid bodies are 
eggs, capable, however, of reproducing the haploid stage if not fertilized; further, 
that the nuclei of the quadrinucleate spores released by diploid individuals are hap- 
loid, and become on germination the nuclei of as many cells of the haploid body. 

The Tilopteridea are believed to represent the evolutionary transition between 
Ectocarpea and the following family. 

Family Dictyotacea [Dictyotaceae] (Hauck) Kjellmann includes about twenty 
genera, Dictyota, Zonaria, Padina, etc., with about one hundred species which are 
commonest on the coasts of warmer oceans. They are thalli of moderate size, erect 
and dichotomously branched or appressed and fan-shaped. They grow by the division 
of a single apical cell or a row of apical cells in each branch. The cells multiplying 
behind the apical cells become differentiated into two tissues, superficial small cells 
rich in plastids and internal larger ones with fewer plastids, forming in different 
species single or multiple layers of cells. 

The Hfe cycle has been studied by Mottier (1898, 1900), Williams (1898), and 
Haupt (1932). There are distinct male haploid individuals, female haploid indivi- 
duals, and diploid individuals, all of the same vegetative structure. The males pro- 
duce sperms from clusters of densely packed plurilocular antheridia. The females 
produce eggs solitary in large oogonia solitary or clustered on the thalli. The eggs 
are without flagella. The diploid individuals produce unilocular sporangia of much 
the same structure as the oogonia. In Zonaria, each sporangium produces eight non- 
motile spores; in Dictyota, each one produces four. 

Order 4. Sporochnoidea [Sporochnoideae] Greville Alg. Brit. 36 (1830). 
Order Chordarieae Greville op. cit. 44. 
Order Chordariaceae Haeckel Gen. Morph. 2: xxxv (1866). 
Orders Desmarestiales and Chordariales Setchell and Gardner in Univ. Califor- 
nia Publ. Bot. 8: 554, 570 (1925). 
Order Sporochnales Sauvageau in Compt. Rend. 182: 364 (1926). 
Brown algae producing motile spores, the haploid stage reduced to scant undiffer- 
entiated filaments, the diploid stage filamentous or thallose, when thallose with apical 
growth. Ralfsia is an exception to the formal characters of the order: it has a haploid 
stage of the same structure as the diploid. This is a rather miscellaneous assemblage, 
rather arbitrarily separated from Phaeozoosporea on the one hand and from Lamin- 
ariea on the other. 

The haploid body of the form of a short-lived body of a few undifferentiated fila- 
ments, like a reduced Ectocarpus, bearing gametangia reduced to single cells, has 

88 ] The Classification of Lower Organisms 

been demonstrated by Kylin ( 1933, 1934, 1937) in a wide variety ot genera, as Asco- 
cyclus, Desmotrichum, Mesogloia, Eudesme, Leathesia, and Stilophora. In the more 
primitive examples, the gametes are not visibly differentiated; in more advanced 
ones, as Carpomitra and Desmarestia, different haploid bodies produce respectively 
smaller sperms and larger eggs, the latter non-motile. 

There is a series of families, Ralfsiacea, Myrionematacea, Myriogloiacea, Meso- 
gloiacea, and others, in which the diploid body consists of filaments differentiated 
into different types. In the simplest of these, the germinating zygote produces in the 
first place a minute thallus-like plate, generally epiphytic on other algae, one cell 
thick, and consisting obviously of branched filaments of limited growth. From this 
plate grow erect filaments. Some of these are simply cylindrical and appear nutritive 
in function; others are attenuate, and may function in protection or in absorbing 
materials from the water; yet others bear the reproductive structures, unilocular or 
plurilocular or both. 

In the more advanced families, the diploid body, after passing through a Ralfsia- 
or Myrionema-Vike stage, may produce a compacted column of filaments with a 
terminal plate of apical cells. Besides adding cells to the column, the apical plate 
gives rise to a fascicle of attenuate hairs projecting forward. Members of the families 
Chordariacea, Sporochnea, and Desmarestiacea produce cylindrical or flattened 
thallose bodies of tridimensionally placed cells differentiated into an outer layer of 
small actively photosynthetic cells and an inner mass of nearly colorless cells. Super- 
ficial hairs, growing in intercalary fashion, may become few, and growth may become 
restricted to a single apical cell. 

By differences in the detailed manner of growth, Setchell and Gardner distin- 
guished two orders among the thalloid forms just mentioned. It is evident, however, 
that the thallose structure (and, likewise, differentiation of gametes) has developed 
repeatedly and independently in the present group. Knowledge which would make it 
possible to divide it into several recognizably natural orders is not yet available. 

Order 5. Cutlerialea [Cutlcriales] Bessey in Univ. Nebraska Studies 7: 289 ( 1907). 

Brown algae producing motile spores, the haploid and diploid bodies being macro- 
scopically visible thalli, alike or different. 

This is a small group, of one family, Cutleriacea, with two genera, Zanardinia 
and Cutlcria, known chiefly from the Mediterranean. In Zanardinia, both haploid 
and diploid bodies are erect and rather freely branched. In Cutlcria, the haploid 
bodies are of this description, while the diploid bodies are appressed and fan-shaped. 
The distinct diploid bodies of Cutlcria were originally named as a different genus, 
Aglaozonia. Falkenberg (1879) first showed that Cutlcria and Aglaozonia arc stages 
of the same thing; Yamanouchi showed that they are respectively a haploid stage 
with 24 chromosomes and a diploid stage with 48. 

The growing margins of the thalli consist of laterally compacted filaments grow- 
ing by the divisions of a band of mcristematic cells which produce free hairs in the 
distal direction and a continuous body of cells in the proximal direction. The latter 
cells are capable of further division, and produce a body several cells thick, with 
small cells rich in plastids on the surface and larger ones with fewer plastids in the 

Haploid individuals bear clusters of stalked plurilocular structures of two types, 
almost always on different individuals, the larger ones consisting of fewer cells which 
release eggs, the smaller of more numerous cells which release sperms. Both kinds of 

Phylum Phaeophyta [ 89 

gametes are flagellum-bearing cells of the type usual in brown algae. The eggs are 
capable of germination without fertilization, reproducing the haploid stage. Diploid 
individuals bear clusters of unilocular sporangia. 

It is only in the life cycle that the Cutlerialea are decidedly different from higher 
Sporochnoidea such as Desmarestia. Their evolutionary origin is explicable by the 
hypothesis of a single mutation which enabled the haploid stage to exhibit the com- 
paratively complicated morphology of the diploid stage, instead of being rudimentary 
as in all Sporochnoidea except Ralfsia (and the exceptional life cycle of Ralfsia 
would be explained by a similar mutation in some primitive example of Sporochnoi- 
dea, such as Myrionema) . 

Older 6. Laminariea [Laminarieae] Greville Alg. Brit. 24 (1830). 

Order Pycnospermeae Kiitzing Phyc. Gen. 333 (1843). 

Order Laminariaceae Haeckel Gen. Morph. 2: xxxv (1866). 

Laminariales Oltmanns Morph. u. Biol. Alg. ed. 2, 2: 2 (1922). 

Order Laminariales Engler and Gilg Syllab. ed. 9 u. 10: 27 ( 1924). 

Order Dictyosiphonales Setchell and Gardner in Univ. California Publ. Bot. 
8: 586 (1925). 

Order Punctariales Kylin in Kungl. Fysiog, Sallsk. Hand!, n. f. 44, no. 7 : 93 
Brown algae with motile spores, the haploid stages reduced to microscopic dimen- 
sions, the diploid stages thallose, growing in intercalary fashion. 

This numerous group, like the preceding small one, is evidently a specialized off- 
shoot from order Sporochnoidea. The familiar examples are the kelps, whose large 
diploid bodies are differentiated into definite members. Kylin considered his order 
Punctariales to represent the transition to the kelps. They are thallose, without dif- 
ferentiation of members, but their microscopic and reproductive characters, as ob- 
served in Soranthera by Angst (1926, 1927), tend to confirm Kylin's opinion, and 
they are accordingly included in the same order with the kelps. Papenfuss (1947) 
pointed it out that the Punctariales of Kylin are essentially the same group as the 
Dictyosiphonales of Setchell and Gardner. 

Sauvageau (1915) first showed that the reproduction of kelps is sexual. The 
grossly visible individuals produce zoospores; these, on germination, produce micro- 
scopic filamentous haploid individuals, generally of distinct sexes, releasing gametes 
from unicellular gametangia. The eggs are without flagella, and it is characteristic 
of them that in emerging from the oogonia they become attached at the opening 
(Kylin, 1916, 1933; Myers, 1928; McKay, 1933; Kanda, 1936; Hollenberg, 1939). 
The same things are true in Soranthera, except that the eggs, although much larger 
than the sperms, are also flagellate. 

The visible bodies of kelps consist of three kinds of members, holdfasts (hapteres), 
being stout root-like growths by which the individuals are anchored to rocks, and 
stalks and blades comparable to stems and leaves. Growth is most active at the sum- 
mits of the stalks. The histology is the same in all members (A. I. Smith, 1939). 
There is a superficial photosynthetic tissue of small cells rich in plastids; on the hold- 
fasts and stalks, this tissue is meristematic, adding cells to the tissue within and in- 
creasing the thickness. Internally there is a cortex of larger cells with fewer plastids. 
In the center there is a medulla containing trumpet fibers, filaments whose cells are 
expanded where they meet and marked by pit-pairs. In the trumpet fibers of Nereo- 
cystis there are actual perforations from cell to cell. The trumpet fibers are not quite 


The Classification of Lower Organisms 

Fig. 16. — Familiar kelps of Pacific North America: a, Egregia Menziesii; h, Nereo- 
cystis Luetkeana; c, Macrocystis pyrifera; d, Postelsia palmaeformis. All approxi- 
mately X /a- 

Phylum Phaeophyta [91 

perfectly analogous to the sieve tubes of higher plants; the nuclei remain alive. The 
minute zoospores are produced in unilocular sporangia. These occur on the surface 
of the body in dense masses, intermingled with, and protected while young by, spe- 
cialized sterile hairs. 

Individuals of Laminaria consist simply of hapteres, a stalk, and one or more 
terminal blades. In various other genera, growth occurs in such fashion as to cause 
the blades to split at the base. With further growth, the splits extend to the margins 
of the blades and increase their number, while intercalary growth at the transitions 
between the stalks and the blades produces elongation and branching of the stalks. 
Early explorers described the stalks of Macrocystis pyrifera as reaching prodigious 
lengths, matters of hundreds of meters, and these accounts have been repeated in 
textbooks down to recent times. Frye, Rigg, and Crandall (1915) found a maximum 
length of somewhat less than fifty meters. The stalks are dichotomously branched 
to a moderate extent and bear series of blades, each with a pear-shaped pneumato- 
cyst or float at the base. The stalks of Nereocystis Luetkeana also were said to be 
extremely long, but the recent observers did not find them to attain fifty meters. They 
are unbranched and bear a single large float from which spring several blades which 
may exceed four meters in length. This great organism is an annual, growing and 
dying within a year. Postelsia palmaeformis, called the sea palm, grows on rocks ex- 
posed to surf. It has erect stalks some 30 cm. tall bearing many pendant linear blades. 
Egregia Menziesii has flattened stalks many meters long with fringes of floats and 
blades along the margins. Laminaria is widely distributed. Macrocystis occurs on the 
northwest coast of North America and in southern oceans. The other kelps which 
have bef:n mentioned are confined to the northwest coast of North America. 

On coasts where they occur, kelps are used as fertilizer. They have been used com- 
mercially as sources of potash, as much as 1-3% of the fresh weight being K as K2O 
(Cameron, 1915); they have been used also as sources of iodine. These uses are not 
economic at most times. 

Setchell and Gardner divided the proper kelps, of which there are about one 
hundred species, into four families. The groups of less elaborate structure which ap- 
pear properly to be placed in the same order are treated by Papenfuss (under Dictyo- 
siphonales) as six families. 

Order 7. Fucoidea [Fucoideae] C. Agardh Syst. Alg. xxxv (1824). 
FucoiDEAE C. Agardh Synops. Alg. Scand. ix (1817). 
Tribe Angiospermeae Kiitzing Phyc. Gen. 349 (1843). 
Order Cyclosporeae Areschoug in Act. Roy. Soc. Upsala 13: 248 (1847). 
Order Fucaceae J. Agardh Sp. Alg. 1 : 180 ( 1848). 
Order Sargassaceae Haeckel Gen. Morph. 2: xxxv (1866). 
Order Fucales Bessey in Univ. Nebraska Studies 7: 290 (1907). 
Order Cyclosporales and suborder Fucineae Taylor in Bot. Gaz. 74: 439 (1922). 
Class Cyclosporeae Kylin in Kungl. Fysiog. Sallsk. Handl. n. f. 44, no. 7: 91 
Thallose brown algae, producing no spores, diploid in all stages except the gametes; 
the latter being sperms, whose posterior flagellum is longer than the anterior one, 
and non-motile eggs. The genus Fucus L. is to be construed as the type genus of order 
Fucoidea, class Melanophycea, and phylum Phaeophyta. 

Two families are usually recognized (others have been segregated). In family 
Fucea [Fuceae] Kiitzing (family Fucaceae Cohn), called the rockweeds, the bodies 


The Classification of Lower Organisms 

are flat dichotomously branching thalli. In family Sargassea [Sargasseae] Kiitzing 
there is a differentiation of holdfasts, stalks, blades, and floats. Growth is by division 
of a single apical cell in each branch or member. There are the usual two tissues, a 
superficial photosynthetic tissue of small cells and an inner tissue of larger cells which 
pull apart to produce a spongy or fibrous mass. 

The gametangia are borne, mixed with sterile hairs, in pits called conceptacles. 
These are clustered, in the Fucea near the tips of branches which have ceased to 
grow (these tips are swollen, and are called receptacles), in the Sargassea on special 
branches. Rarely, oogonia and antheridia occur in the same conceptacles; not infre- 
quently, they occur in different conceptacles on the same individuals; commonly, they 
occur on different individuals. Male and female conceptacles may be distinguished 
by color, the male being orange-yellow, the female of the same dark color as the thalli. 

Male conceptacles are full of branching hairs bearing minute antheridia. In each 
antheridium, the original single nucleus undergoes six successive simultaneous divi- 
sions, producing sixty-four nuclei. These become the nuclei of sperms. Female con- 
ceptacles contain fewer, larger, oogonia, in which the nuclei divide three times, pro- 

FiG. 17. — Microscopic reproductive structures of Laminaria yezoensis after Kanda 
( 1938) : a, male haploid individual releasing sperms; b, sperm; C, zoospore; d, female 
haploid individual of three cells; e, female individual with an egg extnided from the 
oogonium and attached in the mouth f, female individual with two young diploid 
individuals attached at the mouths of oogonia. All x 1,000. 

Phylum Phaeophyta [ 93 

ducing eight. In Fucus, these become the nuclei of as many eggs. In other genera, 
the number of functional eggs is reduced by degeneration of some of them, or of some 
of the nuclei before cell division. In Sargassum, Kunieda (1928) found each oogon- 
ium to produce a single egg in which seven nuclei undergo dissolution while one re- 
mains to function. 

The first two nuclear divisions in each gametangium are meiotic. Farmer and 
Williams (1896) and Strasburger (1897) showed that the bodies are diploid; Yama- 
nouchi (1909) first gave a full account of the meiotic process. The haploid chromo- 
some number of Fucus vesiculosus is 32. In Sargassum Horneri Kunieda found it to 
be 16. 

By a swelling of colloidal material in the conceptacles, the gametangia are forced 
out into the water, where they burst and release the gametes. Fucus was one of the 
first organisms in which syngamy was observed. Thuret (1855) saw multitudes of 
sperms swarm about the eggs, and showed that without sperms the eggs would not 
develop. This much had already been observed in frogs and certain fishes; the dis- 
covery that the essential process is the union of just one sperm with the egg was not 
made until later. The growing zygotes give rise directly to diploid thalli. 
■ The gametangia of the Fucoidea appear to be homologous with the unilocular 
sporangia of other brown algae. In the gametangia, as in unilocular sporangia, the 
meiotic divisions are followed by a few divisions of the haploid nuclei: the Fucoidea 
are not quite perfect examples of the reduction of the haplod stage to the gametes 
only. As to which other brown algae may have provided their evolutionary origin, 
there is no very satisfactory hypothesis; Sporochnus shows certain resemblances. 

Chapter VII 

Phylum 3. PYRRHOPHYTA Pascher 

Order Astoma Siebold in Siebold and Stannius Lehrb. vergl. Anat. 1: 10 (1848). 

Order Phytozoidea Perty Kennt. kleinst. Lebensf. 161 (1852). 

Flagellata Cohn in Zeit. wiss. Zool. 4: 275 (1853). 

Orders Flagellata and Cilio-flagellata Claparede and Lachmann Etudes Infus. 

1: 73 (1858). 
Suborder Mastigophora Diesing in Sitzber. Akad. Wiss. Wein Math. -Nat. CI. 

52, Abt. 1: 294 (1866). 
Stdmme Flagellata and Noctilucae Haeckel Gen. Morph. 2: xxv, xxvi (1866). 
Class Flagellata Kent Man. Inf. 1: 27, 211 (1880). 
Class Mastigophora and orders Flagellata, Dinoflagellata, and Cystoflagellata 

Butschli in Bronn Kl. u. Ord. Thierreichs 1, Abt. 2, Inhalt (1887). 
Class Peridineae Wettstein Handb. syst. Bot. 1: 71 (1901). 
Divisions Flagellatae and Dinoflagellatae Engler Syllab. ed. 3: 6, 8 (1903). 
Pyrrhophyta, Eugleninae, and Chloromonadinae Pascher in Ber. deutschen Bot. 

Gess. 32: 158 (1914). 
Stdmme Pyrrhophyta and Euglenophyta, and Abteilungen Cryptophyceae, Des- 
mokontae, and Dinophyceae, Pascher in Beih. bot. Centralbl. 48, Abt. 2: 325, 
326 (1931). 
Division Pyrrhophyta G. M. Smith Freshw. Algae 10 (1933). 
Protistes trichocystiferes ou progastreades Chadefaud in Ann. Protistol. 5: 323 

Phyla Pyrrhophycophyta and Euglenophycophyta Papenfuss in Bull. Torrey Bot. 

Club 73: 218 (1946). 
Unicellular or colonial organisms, typically with brown or green plastids, flagel- 
late, the flagella solitary or more than one and unequal, the cells marked by grooves 
or pits and sometimes containing trichocysts, i. e., minute structures which lie close 
to the cell membrane and eject thread-like bodies when stimulated. 

The organisms included here are the ones conventionally treated as four orders of 
pigmented flagellates, cryptomonads, dinoflagellates, euglenids, and chloromonads. 
These groups include organisms of the same varied body types, algal, amoeboid, and 
chytrid, that occur in other groups in which the flagellate body type is construed 
as typical. Peridinium may be considered to be the type of the phylum. 

Deflandre (1934) designated as stichoneme [stichonemate) the type of flagcllum 
which bears a single file of appendages, and which had been discovered by Fischer 
(1894) in Euglena. Petersen (1929) reobserved the stichoneme flagellum of Etiglena, 
and found it also in other euglenids, Phacus and Trachelomonas. Deflandre found 
that one flagellum is stichoneme in various further euglenids (but not in all), and 
also in the dinoflagcllate Glenodinium. This is the only report of a stichoneme flagel- 
lum outside of the euglenid group. The fine structure of the flagella of cryptomonads 
and chloromonads has not been determined. 

In some cryptomonads, as Chilomonas, the cells contain granules which stain 
blue with iodine; if these are not starch, one knows not what to call them. Dino- 
flagellates produce a so-called starch which gives a reddish color with iodine, and 
many of them have walls of a so-called cellulose which gives a reddish color with 

Phylum Pyrrhophyta [ 95 

zinc chlor-iodide. The euglenids store granules of a white solid believed not to be 
starch and called paramylum. 

The plastids of cryptomonads and dinoflagellates are of various colors, oflF-color 
green, yellow, brown, bluish, or red. Those of dinoflagellates contain chlorophylls 
a and e; the latter is an exceptional chlorophyll which occurs also in Tribonema. 
Euglenids and chloromonads are typically of the same bright green color as typical 
plants, and the euglenids are known to have the same chlorophylls, a and b, as 
typical plants (Strain, in Franck and Loomis, 1949). 

The groups here brought together exhibit family resemblances in details of the 
mitotic process, so far as these are known. The nuclear membrane usually persists 
through the process. In many examples the chromosomes appear to be present at all 
times, and are quite numerous, elongate, and of the appearance of strings of beads. 
In mitosis, quite as one would assume, they divide lengthwise; the point had been 
disputed, and was established by Hall (1923, 1925, 1937) and Hall and Powell 
(1928). There is a neuromotor apparatus consisting of a centrosome at or near the 
nuclear membrane together with one or more rhizoplasts connecting it to as many 
blepharoplasts at the bases of the flagella. No spindle has been seen, unless the 
peculiar structure, seen in Noctiluca outside of and next to the dividing nucleus, is 
such. The centrosomes may lie at the sides of the dividing nucleus instead of at its 
ends. In the euglenids and some dinoflagellates the nucleus contains a nucleolus-like 
body which does not disappear during mitosis, but divides as the chromosomes do. 

There are few reports of sexual processes in this group. 

Pascher (1914) united the crytomonads and dinoflagellates in a group which 
he named Pyrrhophyta. He and those who follow him leave the euglenids as an iso- 
lated group. Tilden (1933) placed the four groups of flagellates with which we are 
here concerned in division Chrysophyceae, while leaving the Phaeophyceae as a 
distinct division. Her arrangement does not appear to be contrary to nature: the 
cryptomonads are apparently not very far removed from the chrysomonads. The 
different arrangement here maintained, by which the brown algae instead of the 
cryptomonads and so forth are placed in the same phylum with the chrysomonads, is 
believed to have the advantage that that phylum as least is well marked by char- 

Chadefaud (1936) proposed a group consisting of the four groups of flagellates 
here under consideration together with the Infusoria: this on the ground that the 
Infusoria also have deeply indented cells containing trichocysts. He did not give 
to his proposed group a place in the taxonomic system by assigning it to a category 
and giving it a Latin name: he called it by the French common names protistes 
trichocystiferes and progastreades. He suggested two ideas: that if a cell marked 
by a considerable indentation should become divided into many cells forming two 
layers, respectively superficial and against the indentation, the resulting structure 
would be a gastrula; and that the gastrula, and, in fact, the kingdom of animals, 
might have come into existence in this fashion. Perhaps because of novelty, these 
ideas seem far-fetched. So far as it concerns flagellates, Chadefaud's grouping appears 
sound and has been followed in giving limits to the present phylum. 

The phylum is treated as a single class. 

Class MASTSGOPHORA (Diesing) Bu'tschli 

Classes Cryptomonadineae , Rhizocryptineae, Cryptocapsineae, Cryptococcineae, 
Desmomonadineae, Desmocapsineae, Dinoflagellatae, Rhizodininae, Dinocap- 

96 ] The Classification of Lower Organisms 

sineae, Dinococcineae, Dinotrichineae, Euglenineae, and Euglenocapsineae 
Pascher in Beih. bot. Centralbl. 48, Abt. 2: 325, 326 (1931). 
Classes Chloromonadina, Euglenoidina, and Cryptomonadina Hollande in Grasse 

Traite Zool. 1, fasc. 1: 227, 238, 285 (1952). 
Further synonymy as of the name of the phylum. 
Characters of the phylum. 

There are about one thousand known species. Clearly, thirteen classes for their 
accommodation, as proposed by Pascher, are excessive; perhaps one goes too far 
in the other direction in making the entire group a single class. The type of the 
class is the euglenid Astasia. This is true because the family Astasiaea was listed 
first in the earliest appearance of the traditional group Flagellata or Mastigophora 
in due taxonomic form, as order Astoma Siebold, If the euglenids are set apart, 
taking with them the class name Mastigophora, the remaining larger class will be 
called Peridinea [Peridineae] Wettstein. 

The traditional four orders are tenably natural; but that of dinoflagellates includes 
about four-fifths of the species, while the chloromonad group is very inconsiderable. 
The system will be more convenient if the former order is divided into three, and if 
the latter is included in the euglenid order. The resulting five orders are distinguished 
as follows: 

1. Pigmentation if present brown, olive, or the 
like; flagella normally two. 

2. Flagella at the anterior end of the cell, 
not moving in longitudinal and trans- 
verse grooves. 

3. Not walled in the flagellate con- 
dition, flagella not markedly dif- 
ferentiated, or not differentiated 

as anterior and circumferential .Order 1 . Cryptomonadalea. 

3. Usually walled in the flagellate 
condition; flagella respectively an- 
terior and circumferential Order 2. Adiniferidea. 

2. Flagella attached laterally, respectively 
longitudinal and circumferential, moving 
in grooves impressed upon the cells. 
3. Not walled in the flagellate con- 
dition Order 3. Cystoflagellata. 

3. Flagellate cells with a wall usually 

of articulated plates Order 4. Cilioflagellata. 

1. Pigmentation if present typically bright 
green, flagella normally solitary, sometimes 
two or more Order 5. Astoma. 

Order 1. Cryptomonadalea [Cryptomonadales] Engler Syllab. ed. 3: 7 (1903). 
Subclass Cryptomonadineae Engler in Engler and Prantl Nat. Pflanzenfam. 

ITeil, Abt. la: iv (1900). 
Cryptophyceae, including Phaeocapsales and Cryptococcales, Pascher in Ber. 

deutschen bot. Gess. 32: 158 (1914). 
Order Cryptomonadinae Pascher Siisswassei-fl. Deutschland 1: 28 (1914). 
Order Cryptomonadina Doflein Lehrb. Prot. ed. 4: 417 (1916). 

Phylum Pyrrhophyta 


Order Cryptomonadida Calkins Biol. Prot. 265 (1926). 

Orders Cryptocapsales and Cryptococcales Pascher in Beih. bot. Centralbl 48, 
Abt. 2: 325 (1931). 
Solitary (exceptionally colonial) cells, usually with one or two plastids of various 
colors, usually observed in the motile condition, then naked, of dorsiventral (excep- 
tionally isobilateral) symmetry, with two anterior flagella which are not markedly 
differentiated or not respectively anterior and circumferential. 

The resting nucleus contains a karyosome, i. e., a globule which occupies most of 



Fig. 18. — a, Cryptomonas sp. b, Rhodomonas baltica after Kylin ( 1935 ) . c, Chi- 
lomonas Parmecium. d, Cyathomonas sp. e, Sennia sp. f. Vegetative cell, and 
g, zoospore of Paradinium Pouchetii after Chatton (1920). All x 1,000. 

its volume and contains most of the chromatin. Dangeard (1910) and Belar (1916) 
have observed details of mitosis. The numerous chromosomes appear within an 
intact nuclear membrane and form a disk- or drum-shaped figure with its axis at 
right angles to the axis of the cell. No granule more massive than the chromosomes 
persists and divides with them. 

About thirty species are known. They may be treated as five families. 
1. Flagellate cells elongate, with one plane 
of symmetry. 

2. Not parasitic, flagella not markedly dif- 

3. Non-motile in the vegetative con- 
dition Family 1. Cryptococcacea. 

3. Flagellate in the vegetative con- 
dition Family 2. Cryptomonadina. 

98 ] The Classification of Lower Organisms 

3. Amoeboid in the vegetative con- 
dition Family 3. Paramoebida. 

2. Parasitic amoeboid organisms, the flag- 
ella of swimming stages respectively 
anterior and trailing Family 4. Paradinida. 

1. Flagellate cells with two planes of symmetry Family 5. Nephroselmidacea. 

Family 1. Cryptococcacea [Cryptococcaceae] Pascher in Beih. Bot. Centralbl. 48, 
Abt. 2: 325 (1931). YdimWy Phaeocapsaceae West British Freshw. Alg. 48 (1904), 
in part; Phaeocapsa is a chrysomonad. Family Phaeoplakaceae Pascher 1. c. Solitary 
or clustered cells, non-motile in the vegetative condition, reproducing by flagellate 
cells of cryptomonad type. Phaeococcus, Cryptococcus, Phaeoplax. Chrysidella in- 
cludes yellowish cells called zooxanthellae, internally symbiotic in Radiolaria, Rhizo- 
poda, sponges, coelenterates, and rotifers. It is believed that the supposed zoospores 
of various amoeboid organisms are actually flagellate reproductive cells of Chrysi- 
della escaping at certain stages of the life cycles of their hosts. 

Family 2. Cryptomonadina Ehrenberg Infusionsthierchen 38 (1838). Family 
Chilomonadidae Kent Man. Inf. 1: 423 (1880). Family Cryptomonadaceae Engler 
Syllab. ed. 3: 7 (1903). Family Chilomonadaceae Lemmermann 1909. Family 
Cryptomonadidae Poche in Arch. Prot. 30: 159 (1913). Flagellate in the vegetative 
condition, the two flagella not markedly differentiated, springing from the anterior 
end of the cells, usually from the mouth of a pit lined by granules of some sort. 
Cryptomonas and Cryptochrysis have brown or yellow plastids; Chromomonas and 
Cyanomonas have blue ones; Rhodomonas has red ones. Chilomonas is a colorless 
saprophyte familiar in infusions. The colorless Cyathomonas, also from infusions, 
was shown by tJlehla (1911) to be related to Chilomonas. 

Family 3. Paramoebida [Paramoebidae] Poche in Arch. Prot. 30: 173 (1913). 
Schaudinn (1896) discovered the sole known species, Paramoeba Eilhardi, in an 
aquarium of sea water. It is an amoeboid organism with the peculiarity that each 
cell contains beside the nucleus an additional body which divides when the nucleus 
does. The cell may form about itself a shell of debris, and within this may undergo 
division into many cells which escape as pigmented swarmers resembling cells of 

Family 4. Paradinida [Paradinidae] Chatton in Arch Zool. Exp. Gen. 59: 444 
(1920). The sole known species, Paradinium Poucheti, is a parasite in the body 
cavity of copepods. The amoeboid cells are linked together by slender pseudopodia 
so as to form a network. The reproductive cells have a shorter anterior flagellum 
and a longer trailing flagellum. 

Family 5. Nephroselmidacea [Nephroselmidaceac] Pascher Siisswasserfl. Deutsch- 
land 2: 'llO (1913). Family Nephroselmidae Calkins Biol. Prot. 267 (1926). Cells 
isobilateral. Cells disk-shaped, the flagella on the margin: Scnnia. Cells laterally 
extended, bean- or kidney-shaped, the indentation anterior and bearing the flagella: 
Protochrysis, Nephroselmis. 

Order 2. Adiniferidea Kofoid and Swczy in Mem. Univ. California 5: 108 
Suborder Adinida Blitschli in Bronn Kl. u. Ord. Thicrreichs 1: 1001 (1885). 
Suborder Prorocentrinea Poche in Arch. Prot. 30: 160 (1913). 
Desmokontae, including Desmomonadales and Desmocapsales, Pascher in Ber. 
deutschen bot. Gess. 32: 158 (1914). 

Phylum Pyrrhophyta [ 99 

Division Desmokontae; classes Desmomonadineae and Desmocapsineae; and 
orders Desmomonadales, Prorocentrales, and Desmocapsales Pascher in Beih. 
bot. Centralbl. 48, Abt. 2: 325 (1931). 
Suborder Prorocentrina Hall Protozoology 142 (1953). 
Solitary cells, mostly flagellate in the vegetative condition, the flagellate stages 
either naked or bearing a close wall of two valves, with two flagella at the anterior 
end, one extending forward while the other is bent circumferentially and causes the 
cell to whirl while swimming. 

The few known organisms of this group may be treated as a single family. 
Family Adinida Bergh in Morph. Jahrb. 7: 273 (1882). Family Prorocentrinen 
Stein Org. Inf. 3, II Halfte: 8 (1883). Family Prorocentrina Butschli in Bronn Kl. 
u. Ord. Thierreichs 1: 1002 (1885). Family Prorocentraceae Schiitt in Engler and 
Prantl Nat. Pflanzenfam. I Teil, Abt. lb: 6 (1896). Prorocentridae Kofoid in Bull. 
Mas. Comp. Zool. Harvard 50: 164 (1907). Family Prorocentridae Poche in Arch. 
Prot. 30: 160 (1913). Desmocapsa, Haplodiniuni, Desmomastix, Pleuromonas, 
Exuviaella, Prorocentrum; minute brown organisms, mostly marine. 

Order 3. CystoflageUata (Haeckel) Butschli in Bronn Kl. u. Ord. Thierreichs 

1, Abt. 2, Inhalt (1887). 

Tribe [group of families] Gymnodinioidae Poche in Arch. Prot. 30: 161 (1913). 

Classes Rhizodininae, Dinocapsineae, Dinococcineae, and Dinotrichineae; orders 

Gymnodiniales, Rhizodiniales, Dinocapsales, Dinococcales, and Dinotrichales 

Pascher in Beih. bot. Centralbl. 48, Abt. 2 : 326 ( 193 1 ) . 

Suborders Gymnodinina, Dinocapsina, and Dinococcina Hall Protozoology 143, 

147, 149 (1953). 

Haeckel ( 1866) made of Noctiluca alone a phylum under the name of Noctilucae. 

He had the carelessness, as it appears, to publish in the same work the synonymous 

phylar name Myxocystoda as a label in a phylogenetic diagram. In 1873 he used a 

third name, CystoflageUata, and Biitschli took this up; in the text of the Klassen 

und Ordnungen ambiguously as an Unterabtheilung or Ordnung, in the table of 

contents definitely as an order. Allman (1872) had shown that Noctiluca belongs to 

the present group. Biitschli did not agree with this opinion, but it is evidently correct, 

and Haeckel's name becomes the valid one for the order to which Noctiluca belongs 

Typical members of the present order are naked motile cells with brown plastids. 

The two flagella are attached near the equator of the cell. One of them extends in a 

posterior direction, in a groove called the sulcus. The other extends horizontally about 

the cell (generally to the right, in the reversed image seen in the microscope), lying 

in a groove called the girdle. The part of the cell anterior to the girdle is called the 

epicone, the part posterior to it, the hypocone. From the typical structure as thus 

described, there are, as will be seen, many deviations. 

The species, of which more than three hundred are known, may be treated as nine 

1. Relatively unspecialized; having stages freely 
propelled by two flagella, with a single girdle, 
no tentacles, and unspecialized eyespots or 
none; not parasitic; commonly pigmented. 

2. Walled and non-motile in the vegeta- 
tive condition Family 1. Phytodiniacea. 

2. Flagellate in the vegetative condition Family 2. Gymnodiniacea. 

100 ] The Classification of Lower Organisms 

1. Not as above, always without photosynthetic 

2. Amoeboid Family 3. Dinamoebidina. 

2. Flagellate or free-floating. 

3. With multiplied girdles, without 
tentacles or specialized light-sensi- 
tive organelles Family 4. PoLYKRiKroA. 

3. With one girdle or none. 

4. Cells more or less isodiamet- 

5. With prominent light-sen- 
sitive organelles, some- 
times with tentacles Family 5. Pouchetiida. 

5. Without light-sensitive or- 
ganelles, with tentacles. 
6. Not exceptionally 

large Family 6. Protodiniferida. 

6. Reaching exceptional 
sizes, to 1 mm. in di- 
ameter Family 7. Noctilucida. 

4. Cells dome-shaped Family 8. Lepodiscida. 

2. Parasitic Family 9. Blastodinida. 

Family 1 . Phytodiniacea [Phytodiniaceae] Schilling in Pascher Siisswasserfl. 
Deutschland 3: 61 (1913). Family Phytodinidae Calkins Biol. Prot. 277 (1926). 
Dinocapsales, Dinocapsaceae, Dinococcales, Dinotrichales, and Dinotrichaceae 
Pascher in Ber. deutschen bot. Gess. 32: 158 (1914). Orders Dinocapsales, Dino- 
coccales, and Dinotrichales, and families Gloeodiniaceae, Hypnodiniaceae, Dino- 
trichaceae, and Dinocloniaceae Pascher in Beih. bot. Centralbl. 48, Abt. 2: 326 
(1931). Organisms with numerous yellow to brown plastids, walled and non-motile 
in the vegetative condition, reproducing by gymnodinioid zoospores. Some fifty 
species are known; it is only recently that Thompson (1949) has found several of 
these in America. Cells multiplying in a gelatinous matrix: Gloeodinium. Cells 
solitary, dividing into several which escape usually in the flagellate condition; with 
smooth ellipsoid walls: Phytodinium, Stylodinium; anvil-shaped, stalked and with 
two horns: Racihorskya; tetrahedral, with horns at each comer: Tetradinium; with 
a ring of about six horns: Dinastridium. Tending to produce filaments; marine: 
Dinothrix, Dinoclonium. 

Family 2. Gymnodiniacea [Gymnodiniaceae] Schiitt in Engler and Prantl Nat. 
Pflanzenfam. I Teil, Abt. lb: 2 (1896). Subfamily Gymnodinida Bergh in Morph. 
Jahrb. 7: 274 (1882). Gymnodinidae Kofoid in Bull. Mus. Comp. Zool. Harvard 
50: 164 (1907). Family Gymnodiniidae Poche in Arch. Prot. 30: 162 (1913). The 
typical unarmored dinoflagellates, free-swimming, with sulcus and girdle, without 
tentacles or a conspicuous light-sensitive organelle, commonly with photosynthetic 

The genus which is most numerous in species is Gymnodinium Stein. It includes 
both pigmented and non-pigmcnted species, mostly marine, occasional in fresh water, 
the girdles nearly equatorial and forming nearly complete circles. The cells readily 
become encysted, and the cysts may grow to large sizes, reaching diameters of 0.5 mm. 
These cysts have been taken for a distinct genus Pyrocystis. Observed in darkness, 

Phylum Pyrrhophyta [ 101 

the protoplasm in the cysts is seen to become luminous in response to disturbance of 
the medium; they are among the agents of phosphorescence at sea. In Gymnodinium 
Lunula the protoplast of each large globular cyst undergoes division into several 
protoplasts which do not immediately become flagellate; each of them becomes 
crescent-shaped, deposits a cell wall, and is released by dissolution of the wall of the 
parent cyst. In the crescent-shaped cysts, the protoplasts divide into several which 
develop flagella and escape as typical gymnodinioid cells. 

In Hemidinium the girdle forms less than a complete circle; in Amphidinium, the 
girdle is close to the anterior end of the cell; in Gyrodinium, it forms a steep left 
spiral; in Cochliodinium it forms a left spiral of more than one and one half turns. 

Family 3. Dinamoebidina nom. nov. Order Rhizodiniales and family Amoehodi- 
niaceae Pascher (1931), not based on generic names. Non-pigmented amoeboid 
organisms producing crescent-shaped cysts which germinate by releasing gymnodini- 
oid zoospores. Dinamoebidium varians Pascher (1916; originally Dinamoeba, but 
there is an earlier genus of this name, and the author changed it). 

Family 4. Polykrikida [Polykrikidae] Kofoid and Swezy in Mem. Univ. California 
5: 395 (1921). Family Polydinida Butschli (1885), not based on a generic name. 
There is a single genus Polykrikos, of only three known species. They are colorless 
predatory organisms of such a structure as might be produced if a cell of Gymnodi- 
nium were repeatedly to enter upon division and fail to complete it. Each elongate 
cell bears a single extended sulcus and a series of girdles; with each girdle are asso- 
ciated the usual two differentiated flagella. Of nuclei there are usually half as many 
as of girdles. The cells contain structures called nematocysts, whose development 
and structure was studied by Chatton (1914). Each nematocyst consists of a conical 
wall, with a peculiar operculum at the broad end, surrounding a minute cavity 
containing fluid and a coiled thread. Nematocysts are supposed to be homologous 
with trichocysts, and to contribute to protection, or to the capture of prey; the points 
seem not fully established. They occur only in this family and the following. 

Family 5. Pouchetiida [Pouchetiidae] Kofoid and Swezy in Mem. Univ. of Cali- 
fcrnia 5: 414 (1921). Each of the gymnodinioid cells contains a light-sensitive ap- 
paratus, the ocellus, consisting of a pigmented area and of one or more transparent 
globes, of unknown composition, serving as lenses. Most species have nematocysts. 
Protopsis, Pouchetia, etc.; Erythropsis, in warm seas, with a prominent tentacle. 

Family 6. Protodiniferida [Protodiniferidae] Kofoid and Swezy in Mem. Univ. 
California 5:111 (1921). Family Pronoctilucidae Lebour Dinofl. Northern Seas 10 
(1925). Predatory organisms, the cells subglobular, without ocellus or nematocysts, 
but with a tentacle. Pronoctiluca Fabre-Domergue 1889 {Protodinifer Kofoid and 
Swezy 1921); 0.v}'rr/iw Dujardin. 

Description of the neuromotor apparatus and process of division in Oxyrrhis 
marina by Hall (1925) provides part of the authority for, and is in good conformity 
to, the remarks on mitosis included above in the description of the phylum. The 
nucleus contains a prominent internal body (endosome) which does not contain the 
material of the chromosomes and does not disappear during mitosis. A centrosome, 
close outside the nuclear membrane, is connected by two rhizoplasts to blepharo- 
plasts at the bases of the flagella. When a cell is to divide, the centrosome divides; 
the daughter centrosomes do not necessarily lie at the poles of the nucleus where 
the chromosomes assemble. Each daughter centrosome appears to generate one 
rhizoplast, blepharoplast, and flagellum to complete the neuromotor apparatus of a 
eel]. In due course, the endosome, nucleus, and cell undergo constriction. 

102 ] The Classification of Lower Organisms 

Family 7. Noctilucida [Noctilucidae] Kent Man. Inf. 1: 396 (1880). The single 
species Noctiluca scintillans (Mackartney) Kofoid and Swezy (1921; usually known 
as A^. miliaris Suriray ) is a predatory marine organism, the subglobular cells reaching 
dimensions exceeding 1 mm., luminescent when stimulated and accordingly contrib- 
uting to phosphorescence at sea. Each cell is marked by an extensive depression 
representing the sulcus; the girdle is obsolete. A part of the area of the sulcus func- 
tions as a cytostome. A tooth in the sulcus represents the transverse flagellum. Present 
are a longitudinal flagellum, minute in proportion to the cell, and a prominent 

Mitosis in Noctiluca has been studied by Calkins (1899), van Goor (1918), and 
Pratje (1921). Adjacent to the nucleus there is a body of differentiated cytoplasm, 
as large as the nucleus, called by Calkins the attraction sphere. Before mitosis, the 
tentacle and flagellum are absorbed. The attraction sphere becomes elongate and 
its central part becomes converted into fibers. The nucleus becomes appressed to, 
and curved about, the bundle of fibers, and the numerous elongate chromosomes 
assemble against this. The two curved margins of the nucleus draw apart along the 
bundle of fibers, appearing to draw the daughter chromosomes with them. Division is 
completed by constriction of the nucleus and disappearance of the fibers, leaving a 
daughter attraction sphere in association with each daughter nucleus. This peculiar 
mitotic process is probably of no phylogenetic significance, being, like the organism 
in which it occurs, an aberrant by-product of evolution. 

Nuclear division may be followed by division of the cell into two, the entire 
process requiring from twelve to twenty-four hours. Alternatively, the nucleus may 
divide repeatedly, each division requiring from three to four hours; the numerous 
nuclei produced are budded off from the cell in small uniflagellate spores. Ischikawa 
( 1891 ) saw conjugation of pairs of cells, and van Goor stated that this is a preliminary 
to the production of spores; Pratje, on the other hand, could find no evidence of 
conjugation. The spores are believed to give rise by direct growth to cells like the 
original one. 

Family 8. Leptodiscida [Leptodiscidae] Kofoid 1905. Large dome-shaped preda- 
tory marine organisms with small flagella or none. Leptodiscus R. Hertwig (1877) 
was placed by Biitschli in order Cystoflagellata as the sole genus in addition to 
Noctiluca; Craspedotella is a comparatively recent discovery of Kofoid. 

Family 9. Blastodinida [Blastodinidae] Chatton in Arch. Zool. Exp. Gen. 59: 
442 (1920). Ordre Blastodinides Chatton in Compt. Rend. 143: 981 (1906). Fam- 
ilies Apodinidae, Haplozoonidae, Oodinidae, and Syndinidae Chatton op. cit (1920). 
Dinoflagellates which are parasitic chiefly in copepods and tunicates, also in other 
animals and in diatoms. As a general rule, after the parasite has grown to a certain 
size, and a multiplication of nuclei has taken place, a part of the protoplast undergoes 
division to form gymnodinioid zoospores, while the remainder resumes growth in 
the host. Schizodinium, Blastodinium, Apodinium, Chytriodinium, etc. 

Order 4. CiUoflagellata Claparede and Lachman Etudes Inf. 1: 394 (1858). 
Family Peridinaea Ehrcnbcrg Infusionsthierchcn 249 (1838). 
Family Dinifera Bergh in MoVph. Jahrb. 7: 273 (1882). 
Order Dinoflagellata BiitschU in Bronn Kl. u. Ord. Thierreichs 1, Abt. 2: Inhalt 

Subclass Peridiniales Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 

lb: V (1896). 

Phylum Pyrrhophyta [103 

Class Peridineae Wettstein Handb. syst. Bot. 1: 71 (1901). 

Division Dinoflagellata Engler Syllab. ed. 3: 8 (1903). 

Dinophyceae and Dinoflagellatae Pascher in Ber. deutschen bot. Gess. 32: 158 

Order Diniferidea and tribe [group of families] Peridinioidae Kofoid and Swezy 

in Mem. Univ. California 5 : 106, 107 ( 1921 ) . 
Order Dinoflagellida Calkins Biol. Prot. 267 (1926). 
Division Dinophyceae, Class Dinoflagellatae, and order Peridiniales Pascher in 

Beih. bot. Centralbl. 48. Abt. 2: 326(1931). 
Suborder Peridinina Hall Protozoology 144 (1953). 
This order is very close to the preceding; its members are distinguished only by 
the presence, while the cells are in the flagellate condition, of cell walls, consisting in 
most examples of separable plates. The name Cilioflagellata is evidence of an early 
error of observation: the circumferential flagellum was mistaken for a whorl of 
cilia. This name and most of its synonyms were published as applying both to the 
preceding order and this. For almost all of these names the type or obvious standard 
example is Peridinium, with the effect that the names belong to the present order. 
There are about five hundred species, prevalently marine. Five families may be 

Family 1. Peridinaea Ehrenberg Infusionsthierchen 249 (1838). Family Peridin- 
idae Kent Man. Inf. 1: 441 (1880). Family Peridiniaceae Schiitt in Engler and 
Prantl Nat. Pflanzenfam. I Teil, Abt. lb: 9 (1896). Ceratiidae Kofoid in Bull. Mus. 
Comp. Zool. Harvard 50: 164 (1907). The typical dinoflagellates, of numerous 
genera and species. The distinctions among them are largely matters of the detailed 
arrangement of the plates making up the walls. Glenodinium, the plates scarcely 
distinguishable. Peridinium, Goniodoma, Goniaulax, Ceratium, Oxytocum, etc. The 
cells of certain species in various genera are ornamented with prominent horns; in 
Ceratium especially the epitheca is drawn out into one long horn, and the hypotheca 
into one, two, or three. Goniaulax becomes abundant at certain seasons, is eaten by 
shellfish, and renders them poisonous. 

The neuromotor apparatus (much as in Menoidium) and the process of nuclear 
and cell division in Ceratium Hirundinella were described by Entz (1921) and Hall 
(1925). Many nuclei lack the endosome; if present, it disappears during mitosis, as 
does also the nuclear membrane. The daughter centrosomes lie at the sides of the 
blunt-ended mitotic figure. When nuclear division is complete, the protoplast ex- 
pands and then becomes constricted in such fashion that each daughter cell receives 
certain plates of the wall; each daughter cell then secretes the plates which it lacks. 
Zederbauer (1904) reported conjugation in Ceratium. He saw an elongate proto- 
plast with each of its ends covered by a complete cell wall. Dividing cells are of 
quite different appearance. 

Families Ptychodiscida, Cladopyxida, and Amphilothida of Kofoid (1907, the 
names in the feminine; explicitly made families by Poche, 1913) are minor segregates 
from Peridinaea. 

Family 5. Dinophysida (Bergh) Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 
1009 (1885). Subfamily Dinophysida Bergh in Morph. Jahrb. 7: 273 (1882). The 
limits of the plates obscure; girdle near the anterior end; sulcus and girdle bordered 
by prominent flanges. Strictly marine, mostly in warmer oceans. Dinophysis, Oxyphy- 
sis, Amphisolenia, Triposolenia, etc. 


The Classification of Lower Organisms 

Fig. 19, — a, Tetradinium javanicum x 1,000 after Thompson (1949). b, Gytnno- 
dinium striatum x 500 after Kofoid & Swezy (1921). C, Gymnodiniian Lunula, 
flagellate cells forming in a cyst x 500, after Kofoid & Swezy op. cit. d, e, f, Din- 
amoehidium varians; amoeboid vegetative cell, cyst, and production of gymnodinioid 
zoospores x 1,000 after Pascher (1916). g, Noctiluca scintillans x 100 after Allman 
(1872). h, Peridinium cinctum x 1,000. i, Triposolcnia Ambulatrix x 500 after 
Kofoid (1907). j, Amphisolcnia laticincta after Kofoid, op. cit. 

Phylum Pyrrhophyta [ 105 

Order 5. Astoma Siebold in Siebold and Stannius Lehrb. vergl. Anat. 1 : 10 ( 1848) . 
Order Phytozoidea Perty Kennt. kleinst. Lebensf. 161 (1852), in part. 
Order Flagellata Claparede and Lachmann Etudes Inf. 1: 73 (1858), in part. 
Order Flagellato-Eustomata Kent Man. Inf. 1: 36 (1880). 
Suborder Euglenoidina Biitschli in Bronn Kl. u. Ord. Thierreichs 1 : 818 ( 1884). 
Abtheilung (suborder) Chloromonadina Klebs in Zeit. wiss. Zool. 55: 391 

Order Euglenoidina Blochmann Mikr. Tierwelt 1, ed. 2: 50 (1895). 
Subclasses Chloromonadineae and Euglenineae Engler in Engler and Prantl 

Nat. Pflanzenfam. I Teil, Abt. la: v, vi (1900). 
Orders Euglenales and Chloromonadales Engler Syllab. ed. 3: 7 (1903). 
Orders Eugleninae and Chloromonadinae Pascher Siisswasserfl. Deutschland 1 : 

29 (1914). 
Orders Euglenida and Chloromonadida Calkins Biol. Prot. 283, 285 (1926). 
Mostly solitary flagellate cells of fresh water, unwalled and capable of contraction 
and writhing movement; the anterior end of each cell (in the flagellate condition) 
penetrated by a pit, the reservoir or cytopharynx, into which contractile vacuoles 
open; having one flagellum, or two, usually unequal, or more, one flagellum of each 
cell usually being stichoneme; mostly producing a solid storage product, not staining 
blue with iodine, called paramylum. 

Jahn (1946) reviewed this group. He recognized four families, to which one 
more, to include the chloromonads, is to be added. 
1. Producing paramylum. 

2. Flagellum with a swelling near the base, 
usually single but formed of two parts 
which join below the swelling; cells 
mostly pigmented. 

3. Non-motile and walled in the vege- 
tative condition Family 1 . Colaciacea. 

3. Flagellate in the vegetative con- 
dition Family 2. Euglenida. 

2. Flagellum not swollen and usually not 
forked near the base; cells not pig- 

3. Cells without internal rod-shaped 

structures; flagella stichoneme Family 3. Astasiaea. 

3. Cells with internal rod-shaped struc- 
tures; flagella acroneme or simple Family 4. ANisoNEMroA. 

1. Not producing paramylum, storing oil Family 5. Coelomonadina. 

Family 1. Colaciacea [Colaciaceae] Smith Freshw. Alg. 617 (1933). Family 
Colaciidae Jahn in Quart. Rev. Biol. 21: 264 (1946). Euglenoid organisms which 
are walled and non-motile in the vegetative condition. There is a single genus 
Colacium, producing dendroid colonies. 

Family 2. Euglenida Stein Org. Inf. 3, I Halfte: x (1878). Family Euglenina 
Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 820 (1884). Family Euglenaceae 
Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 2: 570 (1897). Solitary 
motile cells, mostly with abundant green plastids, the flagella with swellings near the 
base, mostly solitary and forked below the swelling. Jahn recognized twelve genera. 
Eutreptia has two flagella; Euglenamorpha has three. Members of the latter genus 


The Classification of Lower Organisms 

Fig. 20. — a., Colacium Arbuscula after Stein (1878). b^ Euglena viridis. c, Eu- 
glena Spirogyra. d, Euglena acus. e, Phacus sp. f, Trachelomonas sp. g, Kleb- 
siella alligata after Pascher (1931). All x 1,000. 

Phylum Pyrrhophyta [ 107 

are entozoic in frog tadpoles; some of them are non-pigmented. Three genera having 
the typical single flagella are among the most familiar of flagellates. Euglena has 
fusiform to cylindrical cells freely capable of writhing changes in shape. Phacus has 
flattened cells with a rigid membrane. In Trachelomonas, the protoplast lies loose 
in a rigid lorica which is often ornamented with spines; variations in the form and 
ornamentation of the lorica have made it possible to distinguish a large number of 

There are accounts of mitosis in Euglena by Keuten (1895), Baker (1926), Rat- 
cliffe (1927) and Hall and Jahn (1929). All observers have seen within the nucleus 
a large globule which divides as the nucleus does and appears to guide the separating 
chromosomes. Keuten applied to it the term nucleolo-centrosome; the implications 
of this term are not confidently to be accepted, and the body will better be called by 
the neutral term endosome. RatclifTe's account of mitosis in Euglena Spirogyra is the 
most detailed. It appears that division is initiated when the endosome buds oflE a 
small granule which migrates to a position just within the nuclear membrane and 
divides. The resulting granules may be regarded as centrosomes. The nucleus moves 
forward within the cell and comes into contact with the cell membrane at the bottom 
of the reservoir. Each centrosome appears to generate, just within the cell membrane, 
a granule recognizable as a blepharoplast; the nucleus then withdraws from the cell 
membrane, but the centrosomes remain connected to the blepharoplasts by rhizo- 
plasts. The flagellum, already split at the base, divides throughout its length into two; 
a new flagellum-base grows out from each blepharoplast and becomes fused to one 
of the halves of the old one not far from the base of the latter. Meanwhile, withm 
the intact nuclear membrane, the chromosomes and endosome are dividing. The 
centrosomes are at the sides of the dividing nucleus. No spindle has been recognized. 
Nuclear division is completed by constriction of the membrane. The cell divides by 
constriction which proceeds longitudinally from the anterior end. The centrosomes 
and rhizoplasts disappear, to be replaced during the next division by new ones. 

Hall and Hall and Schoenborn (in several papers, 1938, 1939) have reported 
experiments on nutrition in Euglena. All species are capable of photosynthesis. Some 
of them, surprisingly, have lost the capacity to synthesize amino acids which usually 
accompanies photosynthesis; and there are transitional species in which some in- 
dividuals possess the capacity to make amino acids and others do not, evidently as 
heritable characters. 

Family 3. Astasiaea Ehrenberg Infusionsthierchen 100 (1838). Family Astasiidae 
Kent Man. Inf. 1 : 375 ( 1880) . Family Astasiina Biitschli in Bronn. Kl. u. Ord. Thier- 
reichs 1 : 826 ( 1884) . Family Astasiaceae Senn in Engler and Prantl Nat. Pflanzenfam. 
I Teil, Abt. la: 177 (1900). Colorless organisms. Deflandre found the flagella sticho- 
neme, as to the single flagella of Astasia and Menoidium, and as to one of the two 
flagella of Distigma. Hall and Jahn (1929) found the flagella not swollen near the 
base. The internal rod-shaped structures which characterize the following family are 

Belar (1915) described mitosis in Astasia, and Hall (1923) described it in 
Menoidium. There is a blepharoplast at the base of the flagellum, and some prepara- 
tions show a rhizoplast connecting this to a centrosome immediately outside the 
nuclear membrane. The blepharoplast divides during the early stages of mitosis, and 
the flagellum appears to divide lengthwise. The daughter centrosomes mark the loci 
toward which the dividing chromosomes move. The chromosome number appears to 
be 12. A dividing endosome like that of Euglena is present. 


The Classification of Lower Organisms 

Scytomonas pusilla Stein {Copromonas subtilis Dobell) occurs in the intestines 
of frogs and toads. When cast out with the feces, it exhibits conjugation as a pre- 
liminary to encystment (Dobell, 1908). 

Family 4. Anisonemida [Anisonemidae] Kent Man. Inf. 1: 429 (1880). Families 
Pernamina and Anisonemina Biitschli in Bronn Kl. u. Ord. Thierreichs 1 : 824, 828 
(1884). Family Peranemaceae Senn in Engler and Prantl Nat. Pflanzenfam. I Teil, 
Abt. la: 178 (1900). Family Heteronemidae Calkins Biol. Prot. 285 (1926). Each 
cell of these colorless organisms bears one conspicuous anterior flagellum; most of 
them bear also a less conspicuous trailing flagellum. The trailing flagellum of Pera- 
nema is grown fast to the cell membrane, and is detected only with difficulty (Hall, 

Fig. 21.— a, Menoidium incurvum. b, c. Stages of mitosis in Menoidium incurvum 
X 2,000 after Hall (1923). d, e, Peranema trichophorum. i, Stage of division in 
Peranema trichophorum after Hall (1934). g, Anisoncma truncatum. h, Ento- 
sipon sulcatum, i-m, Vacuolaria viridis: i, cell; j, neuromotor apparatus after Fott 
(1935); k-m, stages of mitosis x 2,000 after Fott, op cit. x 1,000 except as noted. 

Phylum Pyrrhophyta [ 109 

1934). Deflandre was unable to find appendages on the flagella of members of this 
family. As in other members of the order, the flagella spring from a deep anterior pit 
in the cell; in this family, the pit is a functional cytopharynx (Hall, 1933). The cyto- 
plasm of Peranema contains three brief rods, the pharyngeal rods or Staborgane, 
lying near the cytopharynx; their function is unknown. Each cell of Urceolus, of 
Anisonema, and of Heteronema contains a single conspicuous rod extending the length 
of the body. Hall and Powell (1928) and Hall (1934) described the mitotic process 
in Peranema, which is much as in Menoidium. 

Family Coelomonadina Butschli in Bronn Kl. u. Ord. Thierreachs 1 : 819 (1884). 
Family Vacuolariaceae Luther in Bihang Svensk. Vetensk-Akad. Handl. 24, part 3, 
no. 13: 19 (1889). Family Chloromonadaceae Engler Syllab. ed 3: 7 (1903). Family 
Thaumatonemidae Poche in Arch. Prot. 30: 155 (1913). Family Chloromonadidae 
HoUande in Grasse Traite Zool. 1, fasc. 1: 235 (1952); family Thaumatomonadidae 
Hollande op. cit. 686. Unicellular organisms, mostly green, with two diiTerentiated 
flagella springing from a large reservoir, producing globules of oil but no solid storage 
product. Klebs apologized for erecting the grossere Abtheilung Chloromonadina for 
the single genus Vacuolaria, and in fact, this genus differs from other members of the 
present order only in one conspicuous character, the failure to produce paramylum. 
Fott (1935) studied the cytology of Vacuolaria. From the base of each flagellum, a 
rhizoplast extends into the cytoplasm, but fails to come into contact with the nucleus. 
Several granules or swellings, not definitely identifiable as blepharoplasts or centro- 
somes, are distributed along the length of each rhizoplast. In mitosis, which takes 
place within an intact nuclear membrane, the numerous subglobular chromosomes 
form a blunt-ended figure much as in Chilomonas. Genera believed to be allied to 
Vacuolaria include the green flagellate Goniostomum; Chysophaeum Lewis and 
Bryan (1941), a marine organism forming non-motile yellow dendroid colonies of 
m.acroscopic dimensions; and the colorless flagellate Thaumatomastix Lauterborn 
(originally named Thaumatonema, but there is among plants an older genus of this 

Chapter VHI 

Phylum 4. OPISTHOKONTA, phylum novum 

Chytridieae de Bary in Bot. Zeit. 16, Beil. 96 (1858). 

Family Chytridieen de Bary and Woronin (1864). 

Family Chytridiaceae Cohn in Hedwigia 11: 18 (1872). 

Chytridineae Schroter in Engler and Frantl Nat. Pflanzenfam. I Teil, Abt. 1 : 
62 (1892). 

Series (Reihe) Archimycetes (Chytridinae) A. Fischer in Rabenhorst Kryp- 
tog.-Fl. Deutschland 1, Abt. 4: 11 (1892). 

Suborders Chrytidiineae and Monoblepharidineae Engler in Engler and Prantl 
Nat. Pflanzenfam. I Teil, Abt. 1: iii, iv (1897). 

Order Chytridineae Campbell Univ. Textb. Bot. 152 (1902). 

Classes Archimycetae and Monoblepharideae Schaff'ner in Ohio Naturalist 9: 
447,449 (1909). 

Class Archimycetes Gaiimann Vergl. Morph. Pilze 15 (1926). 

Uniflagellatae Sparrow Aq. Phyc. 21 (1943). 

Parasites and saprophytes of simple structure (filamentous, of uniform diameter 
or tapering; or unicellular, with or without rhizoids, i. e. tapering filamentous out- 
growths), with cell walls of chitin, containing no cellulose; producing motile cells 
with solitary posterior acroneme flagella. Type, Chytridium Olla Braun. From 
6tt[o9ioc;, rearward, and KOVT6q^oar. 

Chytrid is the English form of the generic name Chytridium, from Greek )(UTp(<;, 
a jug. Braun (1856) applied this name to a colorless unicellular organism found 
attached to green algae whose cells are penetrated by rhizoids which draw food from 
them and kill them. By chytrids we mean organisms of body types of the general 
nature of that of Chytridium. All such organisms were formerly treated as a single 
taxonomic group. Couch (1938, 1941) showed that the organisms of chytrid body 
type form three markedly distinct groups distinguished by types of flagellation. The 
proper chytrids, those which legitimately constitute a taxonomic group, are marked 
by swimming cells with solitary posterior acroneme flagella, and further by lack of 
cellulose in the cell walls. The group thus marked includes, beside organisms of 
chytrid body type, a few organisms of the filamentous body type of the typical fungi. 

The cytoplasm of members of this group is described as peculiarly lustrous and 
as containing shining globules. In mitosis (seen repeatedly, as by Dangeard, 1900, 
Stevens and Stevens, 1903, Wager, 1913, and Karling, 1937), the sharp-pointed 
spindle forms within the intact nuclear membrane. Some observers have seen centro- 
somes at the poles. The nuclear membrane disappears toward the end of the process. 

The formation of motile cells (zoospores and sometimes gametes) occurs in en- 
larged cells. In these cells there are repeated simultaneous nuclear divisions. After 
the last of these, uninucleate protoplasts, each one containing, ordinarily, one of the 
above-mentioned shining globules, are separated by cleavage. On each of these 
protoplasts a flagellum grows from the cell membrane at the point nearest that part 
of the nucleus which represents a pole of the previous mitotic spindle. Among the 
Blastocladiacea, the nucleus lies against the cell membrane and the flagellum appears 
to spring from a granule within it (Cotner, 1930; Hatch, 1935). Similarly, in Clado- 

Phylum Opisthokonta [111 

chytrium, it appeared to Karling (1937) that the nucleolus generates the flagellum. 
Within the developing swimming cell a body of granules assembles and produces a 
"cap," prominent in stained material, on the anterior side of the nucleus, that is, on 
the side away from the flagellum. 

Nowakowski (1876) observed sexual processes in Polyphagus, and Scherffel 
(1925) observed them in many other chytrids. Sexual processes were known in 
Monoblepharis from the discovery of this genus, and have been studied in detail in 
Allomyces by Emerson (1939, 1941) and Emerson and Wilson (1949). 

The group thus characterized is of fewer than three hundred known species. One 
takes no satisfaction in making it a phylum, but feels constrained to do so by its 
isolation. Note has been taken that other groups including organisms of chytrid 
body type, as Hyphochytrialea, Lagenidialea, and Phytomyxida, have nothing to do 
with the proper chytrids. Furthermore, it will not do to thrust the proper chytrids 
in with the groups of colorless flagellate and amoeboid organisms treated below as 
phylum Protoplasta. One does not trust that group as natural, but it has a morpo- 
logical continuity which would be defaced by the addition of this one. 

Vischer, 1945, coined the name Opistokonten for organisms whose motile cells 
have posterior flagella. Gams (1947) listed as such the green organisms Pedilomonas 
and Chlorochytridion; the choanoflagellates; the proper chytrids; the Sporozoa (the 
whole group by virtue of such examples as have flagellate stages); and the proper 
animals. He inferred that these groups make up a major natural group derived 
from the lowest green algae. This interesting hypothesis must as yet be treated as 
far-fetched. Pedilomonas is scarcely known; it was described by Korschikoff, 1923, 
as a green flagellate of somewhat the appearance of a Chlamydomonas lacking one of 
its flagella. The flagella of the choanoflagellates are pantacroneme instead of acro- 
neme. There remains a striking resemblance between the motile cells of the proper 
chytrids and the sperms of animals. The nuclear cap of the former is quite similar, 
in development and structure, to the beak of the latter. 

The Opisthokonta are reasonably treated as a single class. 

Class ARCHIMYCETES (A. Fischer) SchaflFner 

Synonymy of the phylum. 

Characters of the phylum. 

Previous authors have arranged these organisms in a sequence from strictly uni- 
cellular forms to typically filamentous forms. In the following treatment, this sequence 
is reversed. The course of the evolution of the group is unknown, and it seems reason- 
able to place the body types in the same sequence as among the Oomycetes. The class 
is treated as two orders, Monoblepharidalea, essentially filamentous, and Chytridinea, 
unicellular or producing filaments which taper or are swollen at intervals. 

Order 1. Monoblepharidalea [Monoblepharidales] Sparrow in Mycologia 34: 115 

Suborder Monoblepharidineae Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 

1: iv (1897). 
Blastocladiincae Petersen in Bot. Tidsskr. 29: 357 (1909). 
Order Blastocladiales Sparrow 1. c. 
Opisthokonta whose bodies consist of filaments of uniform diameter, or are of 
types apparently immediately derived from this. Saprophytes in fresh water or soil, 
chiefly on vegetable remains. There are two families. 

112 ] The Classification of Lower Organisms 

Family 1. Monoblepharidacea [Monoblepharidaceae] A. Fischer in Rabenhorst 
Kryptog.-Fl. Deutschland 1, Abt. 4: 378 (1892). Gonapodiineae and Gonapodiaceae 
Petersen in Bot. Tidsskr. 29: 357 (1909). Producing extensive coenocytic filaments, 
non-septate but with false septa of cytoplasm, anchored by rhizoids, reproducing 
asexually by zoospores produced in sporangia which are usually terminal on the 
filaments, the gametes produced in smaller antheridia and larger oogonia which are 
in the more familiar forms terminal and subterminal on the filaments, the branches 
commonly proliferating below them, the eggs without flagella. 

The species, about a dozen, form three genera. In Monoblepharis, the zygote, 
being the entire protoplast of the oogonium, moves out of the oogonium through a 
terminal pore, becomes attached in the opening, and develops a thick wall. In 
Monoblepharella the zygote, retaining the flagellum of the sperm, swims for a time 
before becoming encysted. Gonapodya resembles Monoblepharella (Johns and Ben- 
jamin, 1954). Myrioblepharis Thaxter is believed not to be an organism; it is de- 
scribed as something which might be produced if sporangia of Monoblepharis were 
parasitized by an infusorian. 

Family 2. Blastocladiacea [Blastocladiaceae] Petersen in Bot. Tidsskr. 29: 357 
(1909). Coenocytic filaments, in some examples of a false appearance of septation, 
of the body type of the Rhipidiacea, i. e., differentiated into a basal cell anchored 
by rhizoids and distal branches bearing reproductive structures, sometimes so re- 
duced that the basal cell bears, or is itself, the reproductive structure; the reproduc- 
tive structures including thin-walled zoosporangia, thick-walled resting spores which 
germinate by releasing zoospores, and gametangia; the gametes morphologically 
uniform or larger and smaller, all bearing flagella. 

These organisms are not familiar, although they are readily isolated by baiting 
pond water, or tap water to which soil has been added, with hemp seeds or pieces of 
fruit. There are four genera, Allomyces, Blastocladia, Blastocladiella, and Sphaero- 
cladia, with about twenty-five known species. Allomyces is of interest for varied life 
cycles, and Blastocladia for a peculiar type of metabolism. 

The first known species of Allomyces, A. Arbuscula, was discovered by Butler 
(1911) on dead flies in water in India. The individuals are of the appearance of 
minuscule shrubs, the branches divided by pseudosepta punctured in the middle and 
ending in series of varicolored reproductive structures. Ordinary sporangia are 
colorless, resting spores are brown, mature antheridia are pink, and mature oogonia 
dull gray. Kniep (1929), in discovering the second species, A. javanicus, found that 
the individuals are of two types, one bearing sporangia and resting spores, the other 
oogonia and antheridia. Thus this organism has a complete life cycle of morpholo- 
gically homologous haploid and diploid individuals. Kniep supposed that meiosis 
occurs in the resting spores, and Emerson and Wilson (1949) established the point. 
The chromosome number (n) oi A. Arbuscula is 7; that of A. javanicus var. macro- 
gynus and of A. cystogenes is 14. 

The life cycle of A. Arbuscula is the same as that of A. javanicus. In A. cystogenes, 
the haploid stage consists merely of the zoospores from the resting spores; these 
become encysted and germinate by releasing isogametes. Thus this species has a life 
cycle essentially of the advanced type characteristic of animals. There are further 
species of Allomyces in which a sexual cycle is believed not to occur. 

In Blastocladia the basal cell bears directly multiple reproductive structures. 
Organisms of this genus are less easily cultured than Allomyces; they require several 
vitamins of the B group (Cantino, 1948). They tolerate oxygen, but do not require it. 

Phylum Opisthokonta [113 

They convert sugars to lactic and succinic acids, producing no CO2; the acids, if not 
neutralized, check the growth of cultures (Emerson and Cantino, 1948; Cantino, 
1949). Blastocladia appears to have lost the capacity to carry on the aerobic stages of 
energesis, thus reverting to the type of metabolism characteristic of the supposedly 
most primitive bacteria. 

In Blastocladiella, the basal cell bears a single reproductive structure. DiflFerent 
species have the same three types of life cycle which occur in Allomyces (Couch and 
WhiflFen, 1942). In Sphacrodadia the vegetative body is reduced to the unicellular 
condition which is characteristic of the following order rather than of this. The life 
cycle is of the complete homologous type. 

Order 2. Chytridinea [Chytridineae] (Schroter) Campbell Univ. Textb. Bot. 

152 (1902). 
Orders Myxochytridinae and My cocky tridinae A. Fischer in Rabenhorst Kryp- 
tog. Fl. Deutschland 1, Abt. 4: 20, 72 (1892), not based on generic names. 
Order Chytridiales Auctt. 

Further synonymy as of the name of the phylum. 

Opisthokonta which consist entirely or largely of more or less isodiametric bodies 
called centers: the centers may send out filaments more slender than themselves, 
generating at their ends further centers; or may be capable only of producing rhizoids, 
i. e., tapering absorptive filaments; or may be by themselves complete individuals. 

The chytrids are commonly thought of as prevalently parasitic on algae and 
higher plants. They attack also rotifers, insects, nematodes, and other minute animals; 
some parasitize other chytrids (Karling, 1942, 1948). It is probable, however, that 
the majority of the group are saprophytic on organic remains. Some have been 
cultured with no other organic food than cellulose (Haskins, 1939); new forms 
have been discovered by baiting with, and culturing on, chitin (Karling, 1945; 
Hanson, 1946) or keratin (Karling, 1946, 1947). 

The following varieties of vegetative structure may be noted, (a) A zoospore, 
settling upon the surface of an appropriate host or substratum, may penetrate this 
by means of a walled filament which develops a terminal center; the center then 
sends out rhizoids, and also filaments which generate further centers, (b) Develop- 
ment may be as above except that only one center is formed. The body thus described 
is of the Entophlyctis type of Sparrow (1943). (c) The zoospore may itself become 
the single center, penetrating its host or substratum only by rhizoids. The resulting 
body is of the Chytridium type if the center is in contact with the host or substratum, 
of the Rhizidium type if it is not. (d) The protoplast of the zoospore may migrate 
into the protoplast of the host and there become a center without rhizoids; the 
resulting body is of the Olpidium type. To the varied bodies thus described, the 
following terminology is applicable: 

Pluricentric, with more centers than one; mono centric, with a single center. 

Intramatrical, the center developing within the substratum or host; alternatively, 
in a host, endobiotic. 

Extramatrical or epibiotic, contrary to the foregoing. 

Eucarpic, the center not constituting the entire body; holocarpic, the center con- 
stituting the entire body. 

The center regularly remains uninucleate during the vegetative phases and then 
becomes the seat of successive simultaneous nuclear division, of cleavage, and of the 
maturation of zoospores. Thus it is converted into a sporangium. In many forms, the 


The Classification of Lower Organisms 

Fig. 22. — Monoblepharidalea: a-f, M o noble pharella Taylori x 1,000 after 
Springer (1945); a, germinating spore producing a filament and a rhizoid; b, spor- 
angium releasing spores; c, empty antheridium and sperm uniting with egg; d, sperms 
escaping from antheridium and zygote escaping from oogonium; e, swimming zygote; 
f. encysted zygote, g-i, Allomyces javanicus x 100 after Kniep (1929); g, asexual 

(Continued bottom p. 115) 

Phylum Opisthokonta [115 

proximal part of the system of rhizoids develops a large swelling called the apo- 
physis. In other forms, the center generates the sporangium as an outgrowth. In 
these circumstances, the center is sometimes called an apophysis, but were better 
called a presporangium. The sporangium discharges its spores, usually, through one 
or more tubes which grow forth from it. The tube may open through a difTerentiated 
cap, the operculum; the production of opercula appears to mark a natural subordi- 
nate group. 

Syngamy occurs in different chytrids in most of the possible fashions, by union of 
like or unlike swimming cells, by the union of a swimming cell with a stationary one, 
or by the establishment of contact by growth. The zygote regularly becomes a thick- 
walled resting spore (asexual resting spores are also of frequent occurrence). Resting 
spores germinate by producing zoospores. Meiosis has not been observed, but is be- 
lieved to occur during the first nuclear divisions in the germinating zygote; the life 
cycle is apparently of the primitive type, in which all cells except the zygote are 
haploid {Phy so derma, or at least some of its species, is believed to be exceptional). 
Sparrow (1943) recognized nine families. One of these does not appear tenable; 
the remainder are distinguished as follows: 
1. Sporangia not opening through opercula. 
2. Eucarpic, i. e., producing rhizoids and 
sometimes other filaments, the centers 
not constituting the entire body. 

3. Pluricentric Family 1. Cladochytriacea. 

3. Monocentric. 

4. Germinating spores generat- 
ing the center as a distinct 

body Family 2. Phlyctidiacea. 

4. Zoospores themselves becom- 
ing centers, and subsequently 

sporangia or presporangia Family 3. Rhizidiacea. 

2. Holocarpic, i. e., without rhizoids, the 
individual consisting entirely of one or 
more centers. 

3. Centers becoming presporangia, 
each one generating a cluster of 

sporangia Family 4. Synchytriacea. 

3. Centers proliferating, giving rise to 

linear series of sporangia Family 5. Achlyogetonacea. 

3. Each center becoming one spor- 
angium Family 6. Olpidiacea. 

individual with light sporangia and dark resting cells with pitted walls; h, branch of 
sexual individual, the oogonia larger and darker than the antheridia; i, gametes. 
j-m, Allomyces Arbuscula after Hatch (1935); j, k, gametes, x 1,000; 1, m, mitotic 
figures in the gametangia, x 2,000. n-r, Blastocladiella cystogena, x 500, after Couch 
and WhifFen (1942); n, individual producing a resting spore; O, resting spore germ- 
inating by release of numerous naked protoplasts; these become flagellate zoospores, 
p, which subsequently encyst; q, the protoplast of each cyst divides to produce four 
gametes; r, young zygote with the flagella of both gametes. 


The Classification of Lower Organisms 

' B^^ 

fiG. 23. — Chytridinea: a-c, Polyphagus Euglenae attacking cells of Euglena, 
X 400, after Nowakowski (1876); in figure b, two individuals have made contact 
and a zygote is developing at the point of junction; c, sporangium, d-i, Olpidium 
Allomycetos attacking Alomyces anomalus, x 1,000, after Karling (1948); d, e, zoo- 
spores; f, sporangium of the host beset with many parasites; g, h, resting cells of the 
host containing respectively sporangia and resting cells of the parasite; i, germina- 
tion of resting cell. 

Phylum Opisthokonta [117 

1. Sporangia opening through opercula. 

2. Pluricentric Family 7. Nowakowskiellacea. 

2. Monocentric Family 8. Chytridiacea. 

Family 1. Cladochytriacea [Cladochytriaceae] Schroter in Engler and Prantl Nat. 
Pflanzenfam. I Teil, Abt. 1: 80 (1892). Family Hyphochytriaceae [Cladochytria- 
ceae) A. Fischer in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 4: 131 (1892), in 
part. Family Physodermataceae Sparrow Aq. Phyc. 304 (1943). Pluricentric chytrids, 
the sporangia not operculate. The members of this family are of the same body type 
(designated by Karling, 1931, the rhizo mycelium) as the anisochytrid Hyphochy- 
trium and the Nowakowskiellacea of the present order. In most Cladochytriacea the 
rhizomycelium includes pairs of swollen cells ("turbinate organs") which give a false 
appearance of conjugation. There are some forty known species, mostly of two 
genera, Cladochytrium, saprophytic in vegetable remains, and Physoderma (including 
Urophlyctis), parasitic in higher plants. Sparrow (1946, 1947) discovered in certain 
species of Physoderma an alternation of morphologically distinguishable generations, 
both on the same hosts; the generations are presumably haploid and diploid, but this 
has not been established by observation of syngamy and meiosis. Polychytrium grows 
well only on chitin (Ajello, 1948). 

Family 2. Phlyctidiacea [Phlyctidiaceae] Sparrow in Mycologia 34: 114 (1942). 
Family Sporochytriaceae [Rhizidiaceae, Polyphagaceae) subfamily Metasporeae A. 
Fischer in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 4: 85 (1892). Monocentric 
eucarpic chytrids, the centers developed at the ends of filaments which grow from the 
zoospores, sporangia without opercula. 

These are the most familiar chytrids. There are more than one hundred species. 
Many are parasitic, on blue-green and green algae, diatoms, pollen grains, nematodes, 
and other minute fresh-water life; others are saprophytic, on cellulose, chitin, or 
keratin. Rhizophidium, the most numerous genus; Phlyctidium, Phlyctorhiza, Ento- 
phlyctis, Diplophlyctis, Loborhiza, etc. 

Family 3. Rhizidiacea [Rhizidiaceae] Schroter in Engler and Prantl Nat. Pflanzen- 
fam. I Teil, Abt. 1: 75 (1892). Family Sporochytriaceae [Rhizidiaceae, Polyphaga- 
ceae) A. Fischer in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 4: 85 (1892) 
and subfamily Orthosporeae op. cit. 124. Monocentric eucarpic chytrids, the zoospores 
enlarging and becoming centers, which in turn become sporangia or presporangia; the 
sporangia without opercula. A moderate number of species, parasitic on blue-green 
or green algae, flagellates, or diatoms; or chitinophilous, saprophytic in the shed 
exoskeletons of insects. Rhizidium, Siphonaria, Asterophlyctis, Polyphagus, etc. 
Polyphargus Euglenae Nowakowski (1876) is a classic example. The centers lie free 
in the water, parasitizing cysts of Euglena through freely branching and widely 
spreading rhizoids. Most centers act as presporangia. Syngamy occurs when a rhizoid 
from one center makes contact with another center. The protoplasm of the latter 
migrates into the tip of the rhizoid, which swells and becomes a resting spore. 

Family 4. Synchytriacea [Synchytriaceae] Schroter op. cit. 71. Family Merol- 
pidiaceae [Synchytriaceae) A. Fischer op. cit. 45. Holocarpic chytrids, the intra- 
matrical cell unwalled in the vegetative condition, becoming a presporangium or a 
resting spore, either of which gives rise to a cluster of sporangia. Synchytrium, 
parasitic on higher plants; Micromycopsis on Conjugatae. 

Family 5. Achlyogetonacea [Achlyogetonaceae] Sparrow in Mycologia 34: 114 
(1942). Chytrids without rhizoids, the intramatrical center proliferating and pro- 
ducing a linear series of centers, each of which becomes a sporangium without an 

118] The Classification of Lower Organisms 

operculum. Achlyogeton, in green algae, diatoms, and nematodes; of very much the 
appearance of certain Lagenidialea. 

Family 6. Olpidiacea [Olpidiaceae] Schroter op. cit. 67. Family Monolpidiaceae 
[Olpidiaceae) A. Fischer op. cit. 20. Holocarpic chytrids, each individual a single 
intramatrical parasitic center, naked until the reproductive phase, when it becomes 
a sporangium without an operculum. Olpidium, attacking blue-green and green algae, 
diatoms, flagellates, Allomyces, Vampyrella, rotifers, and nematodes. Rozella, attack- 
ing Oomycetes and producing spiny resting spores, has been confused with certain 
Lagenidialea. The genera Sphaerita and Nucleophaga of Dangeard, including 
intracellular parasites of amoebas and Infusoria, have been placed in this family; 
it seems more probable that they should be placed among bacteria of family Rickett- 

Family 7. Nowakowskiellacea [Nowakowskiellaceae] Sparrow in Mycologia 34: 
115 (1942). Family Megachytriaceae Sparrow Aq. Phyc. 378 (1943). Pluricentric 
chytrids, the sporangia with opercula. A moderate number of saprophytes on material 
of green algae and higher plants. Nowakowskiella, Megachytrium, etc. Zygochytrium 
was described by Sorokin, 1874, as living on decaying insects, producing multiple 
operculate sporangia, and exhibiting a conjugation of filaments to produce zygotes 
much like those of Zygomycetes. It has apparently not been reobserved. 

Family 8. Chytridiacea [Chytridiaceae] Cohn in Hedwigia 11: 18 (1872). Family 
Chytridieen de Bary and Woronin in Berichte Verhandl. Naturf. Gess. Freiburg 3 
(Heft 2) : 46 ( 1864). Monocentric eucarpic chytrids, the sporangia operculate. Some 
fifty species, the majority parasitic on fresh water algae. Chytridium, etc. Catenochy- 
tridium, saprophytic in cast-off exoskeletons of insects. 

Chapter IX 

Phylum 5. INOPHYTA Haeckel 

Order Fungi L. Sp. PI. 1 1 7 1 (1 753 ) . 

Hysterophyta Link, 1808. 

Classes Fungi and Lichencs Bartling Ord. Nat. 4 (1830). 

Regnum Mycetoideum Fries Syst. Myc. 1: Ivi (1832). 

Class Lichenes and section Hysterophyta with class Fungi Endlicher Gen. PI. 11, 
16 (1836). 

Stamm Inophyta Haeckel Gen. Morph. 2: xxxvi (1866). 

Subdivision Fungi Engler and Prantl Nat. Pflanzenfam. II Teil: 1 (1889). 

Division Eumycetes Engler Syllab. ed. 3: 25 (1903). 

Phylum Carpomyceteae Bessey in Univ. Nebraska Studies 7: 249 (1907). 

Stamm Mycophyta Pascher in Beih. bot. Centralbl. 48, Abt. 2: 330 ( 1931 ). 

Kingdom Mycetalia Conard Plants of Iowa iv (1939). 

Phylum Eumycophyta Tippo in Chron. Bot. 7: 205 (1942). 

Parasites and saprophytes without flagellate stages, the bodies filamentous, the 
w;,lls containing no cellulose. 

This group represents the conventional division or subdivision Fungi of the 
kingdom of plants, excluding, of course, the bacteria, Oomycetes, chytrids, and 
Mycetozoa. The name Fungi, used as a scientific name, is properly to be applied, 
by authority of Linnaeus, to an order. Agaricus campestris L. will be recognized 
as the standard species of the phylum and of the order. 

Those who study Inophyta are accustomed to use, for soma and filament respec- 
tively, the terms mycelium and hypha. The walls of the hyphae are believed to consist 
of pectic material. A small percentage of chitin is usually present (Schmidt, 1936); 
cellulose is totally absent (Thomas, 1928; Nabel, 1939; Castle, 1945). The organism 
Basidiobolus, having hyphae walled with cellulose, is tentatively retained among 
Inophyta as an exception. 

The multiplication and dissemination of those organisms is by spores, of various 
types, scattered in the air. Most Inophyta produce two or more kinds of spores, some 
of them asexually, others as features of a sexual cycle. Spores produced within cases 
are called endospores, and the cases sporangia. Other spores are produced externally, 
commonly by constriction of the ends of hyphae. Spores thus produced are called 
conidia, and the hyphae or other structures which bear them, conidiophores. Spores 
are commonly produced not directly on the mycelium but on macroscopic structures 
of various types, all of which may be called by the familiar term fruit. The common 
mushroom as we see it is a fruit; it is the temporary spore-producing structure of 
an organism whose soma consists of filaments living saprophytically in the soil 

It is expedient to mention at this point the growths called lichens, which are 
traditionally treated as a taxonomic group, either subordinate to Fungi or of the 
same rank. Lichens are gelatinous or thallose growths, usually of an impure green 
color, common everywhere, terrestrial or epiphytic, as on stones, trees, or fence 
posts. The microscope, in the hands of de Bary and others, showed that they consist 
of cells of two types, colorless filaments like those of Inophyta, and pigmented 

120] The Classification of Lower Organisms 

cells of quite the character of those of certain algae. De Bary (in Hofmeister, 1866) 
concluded that some lichens are not organisms but combinations of totally diverse 
organisms. Presently (1868) he was convinced by the work of Schwendener, soon 
(1868) published under his own name, ". . . dass die Flechten sammt und senders 
keine selbststiindigen Pflanzen seien, sondern Pilze aus der Abtheilung der Ascomy- 
ceten, denen die fraglichen Algen — deren Selbststandigkeit ich also nicht bezweifle — 
ah Nahrpflanzen dienen." In 1879 de Bary coined the term symbiosis to designate 
the association of different kinds of organisms. In de Bary's usage the term included 
parasitism; in general usage, it means association to mutual advantage. The lichens 
are a classic example of symbiosis. 

Clearly, the group of lichens is not to be maintained; the algal components are 
known to have natural places among algae, and the inophyte components are to be 
assigned to their natural places among Inophyta, almost all in various orders of 
class Ascomycetes. This has already been done by Clements (1909) and Clements 
and Shear ( 1931 ). The numerous names which students of lichens have given to them 
are to be applied to the inophyte components. 

Another common example of symbiosis involving inophytes is furnished by at least 
some of those which live on or in the tissues of higher plants without killing them 
(Kelley, 1950). They occur mostly on roots. Frank (1885) coined the term mycorhiza 
to designate the combination of roots and inophytes; it will be more convenient to 
hold that this term designates the inophyte component of the combination. Such 
mycorhizae as cover the growing tips of roots are helpful to their hosts by serving as 
agents of absorption. 

Jones (1951) estimated the number of species of Inophyta as 40,000. This is 
surely an extreme underestimate. Martin (1951) gives reason for believing the num- 
ber to be about as great as that of flowering plants, of the order of 300,000. 

The early classifications of "fungi," as by Persoon (1801) and Fries (1821-1832), 
were based on gross characters. They presented, along with recognizable groups 
whose names are to be applied in order of priority, others which were mere random 
assemblages, and whose names are to be abandoned as nomina confusa. De Bary (in 
Hofmeister, 1866; 1884), having applied comparatively modern methods, established 
a dozen groups (under German names). These, so far as they are retained in the 
present phylum, have been assembled as three classes distinguished by details of the 
sexual cycle. A fourth class, acknowledgedly artificial, is maintained for the accomo- 
dation of the numerous and important fungi whose sexual cycles are unknown. The 
termination -mycetes, of the names of the classes and also of various subordinate 
groups, is the Greek ^uKr]T£q, the plural of (auKT^c;, a mold or mildew. The termi- 
nation -mycetae which some authors have used is a solecism. 

1. Reproducing sexually, or by apomictic pro- 
cesses clearly of sexual origin. 
2. The zygote becoming a thick-walled 

resting cell; fruits none or inconsiderable Class 1. Zygomycetes. 

2. The zygote not becoming a thick-walled 
resting cell; mostly producing fruits. 
3. The zygotes giving rise, usually in- 
directly, to sporangia called asci, 
each typically containing eight 
spores called ascospores Class 2. Ascomycetes. 

Phylum Inophyta [121 

3. The zygotes giving rise indirectly to 
conidiophores called basidia, each 
bearing typically four conidia 

called basidiospores Class 4. Basidiomycetes. 

1. Not known to reproduce sexually Class 3. Hyphomycetes. 

Class 1. ZYGOMYCETES (Sachs ex Bennett and Thistleton-Dyer) 


Zygomyceten Sachs Lehrb. Bot. ed. 4: 248 (1874). 

Zygomycetes Bennett and Thistleton-Dyer in Sachs Textb. Bot. English ed. 847 

Class Zygomycetes Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 1: 
32 (1879). 

Order Zygomycetes Engler Syllab. 23 (1892). 

Class Zygomyceteae Schaffner in Ohio Naturalist 9: 449 ( 1909). 

Inophyta whose zygotes are thick-walled resting cells, in germination giving rise to 
spores indistinguishable from those produced asexually; hyphae usually without cross- 
walls; mostly not producing fruits. The standard species is Mucor Mucedo L. 

Among the Inophyta as here limited, the Zygomycetes appear to be primitive (an 
alternative hypothesis, that certain Ascomycetes are primitive, will be discussed be- 
low). Traditionally, the Zygomycetes are associated with the Oomycetes. The asso- 
ciation is probably mistaken, being based merely on similarity of body form: the 
Zygomycetes are terrestrial instead of aquatic, produce no flagellate cells, have no 
cellulose in their cell walls (except in Basidiobolus) , and do not produce female 
gametes by the cutting out of cells within a cell. In later editions of Engler's Syllabus 
(1924), one finds most of the chytrids included among the Zygomycetes, instead of 
in their conventional place among the Oomycetes. The hypothesis thus suggested, 
that the Opisthokonta may represent the ancestry of the Inophyta, is attractive, but 
not to present knowledge supported by convincing evidence. Class Zygomycetes and 
phylum Inophyta must as yet be regarded as of unknown origin and treated as isolated. 

There are some 500 known species of Zygomycetes. They form two orders. The 
bulk of the group, and the typical examples, are order Mucorina. A minority, 
distinguished by parasitism and by explosively discharged conidia, are order 

Order 1. Mucorina [Mucorini] Fries Syst. Myc. 3: 296 (1832). 

Suborder Mucorineae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, 

Abt. 1: iv (1897). 
Order Mucorineae Campbell Univ. Textb. Bot. 158 (1902). 
Order Spirogyrales (presumably in part only) Clements Gen. Fung. 12 (1909). 
Order Mucorales Smith Crypt. Bot. 1 : 405 (1938). 
Order Zoopagales Bessey Morph. and Tax. Fungi 117 (1950). 
The typical Zygomycetes, mostly saprophytic, not producing explosively dis- 
charged conidia [Piloholus produces explosively discharged sporangia). 

The asexual reproductive structures of the supposedly primitive Mucorina, as 
Mucor and Rhizopus, are solitary globular sporangia terminal on erect hyphae. In 
the developing sporangium, a dome-shaped basal sterile area, the columella, is set 
apart by cleavage followed by deposition of a wall. The protoplasm above the 


'I'hc Classification of Lower Organisms 

Fig. 24. — Zygomycetes: a-d, Rhizopus nigricans; a, sporangia x 50; b-d, prega- 
mctes, suspensors and gametes, and zygote x 200. e. Zygote of Phycomyccs nitens 
after Blakeslcc ( 1904). f, g, Conidiophore with young conidia, and mature conidia, 
of Syncephalis pycnosperma after Thaxtcr (1897). h, i, Conjugation of Synce- 
phalis nodosa after Thaxter, op. cit. j, Sporangium of Synccphalastrum raccmosum 
after Thaxter, op. cit. k, Sporangium of Flaplospoiangium lignicola after Martin 
(1937), x 1,000. 

Phylum Inophyta [ 123 

columella undergoes cleavage to form spores, which may remain plurinucleate 
(Swingle, 1903). Other members of the order exhibit transitions (apparently two 
distinct series of transitions) from sporangia as just described to typical conidia. 

Syngamy occurs when the tips of pairs of hyphae meet and are cut off by crosswalls 
to act as multinucleate gametes. The process is regarded as conjugation, although 
the gametes of a pair are usually not of the same size. Conjugation does not occur 
at random, but, in most Zygomycetes, between branches from hyphae of two mating 
types, designated plus and minus (the distinction of mating types is not identical 
with the differentiation of sexes). Zygomycetes were the first group reproducing by 
conjugation in which a distinction of mating types was discovered; the discovery was 
by Blakeslee (1904). 

Syngamy is preceded by a flare of mitoses in the gametes. The mitotic figures are 
sharp-pointed, as though centrosomes were present; the haploid chromosome number 
appears to be 2. The process is not meiotic (Moreau, 1913). After these divisions, 
the walls between the gametes break down and the nuclei unite in pairs. Unpaired 
nuclei, presumably contributed in excess by one gamete or the other, undergo disso- 
lution (Keene, 1914, 1919). Ordinarily, the zygote enlarges and becomes a thick- 
walled resting spore; in some examples, the resting spore forms as an outgrowth on 
what was one of the gametes. In Phycomyces, Absidia, and Syncephalis, the hyphae 
which have produced the gametes, and to which the zygote remains attached, .send 
out branches which form a layer about the zygote. These branches might be inter- 
preted as making up fruits. Endogone produces definite fruits of considerable size. 

A zygote germinates by production of a hypha bearing a sporangium (Blakeslee, 
1906). Meiosis is believed to occur in the course of germination. 

While Mucorina in general are saprophytic, some of them are parasitic on others, 
Piptocephalis and Chaetocladhim on Mucor, and Parasitella on Absidia. Drechsler 
(1935, 1937) discovered a number of organisms apparently of this group parasitizing 
amoebas and nematodes in the soil. 

The Mucorina may be treated as five families. 
1. Not producing macroscopic fruits. 

2. Not parasitic on amoebas or nematodes. 
3. All spores produced in sporangia 

with columellae Family 1. Mucoracea. 

3. Not as above. 

4. Producing sporangia or else 
conidia as outgrowths from a 
knob, homologous with a 
sporangium, solitary on an un- 

branched stalk Family 2. Piptocephalidacea. 

4. Sporangia or conidia solitary 
and terminal on branches of a 
branched sporangiophore or 
conidiophore; sporangia, if 

produced, without columellae Family 3. Mortierellacea. 

2. Parasitic on amoebas or nematodes Family 4. Zoopagacea. 

1. Producing macroscopic fruits Family 5. Endogonacea. 

Family 1. Mucoracea [Mucoraceae] Cohn in Hedwigia 11: 17 (1872). Mucorina 
whose spores are produced exclusively in sporangia with columellae solitary on un- 
branched sporangiophores. Mucor L., typified by M. Mucedo, is now limited to a 

124] The Classification of Lower Organisms 

small group mostly saprophytic on manure. Pilobolus, another coprophilous genus, 
is distinguished by sporangiophores which become swollen at the summit, bend 
toward the light, and discharge the sporangia violently to a distance of several meters. 
Rhizopus nigricans, the common black bread mold; Phycomyces, Ahsidia, Sporodinia, 

Family 2. Piptocephalidacea [Piptocephalidaceae] Schroter in Engler and Prantl 
Nat. Pflanzenfam. I Teil, Abt. 1: 132 (1893). Family Choanephoraceae Schroter op. 
cit. 131. Mucorina producing sporangia without columellae, or conidia, in compact 
clusters terminal on unbranched stalks. Blakesleea, transitional between the preceding 
family and this, may produce solitary sporangia with columellae, or else, as out- 
growths from the primordia of sporangia, clusters of minuscule sporangia without 
columellae. Cunninghamella, producing heads of globular conidia; Syncephalastrum, 
with clustered cylindrical sporangia; Syncephalis and Piptocephalis, producuig 
clustered chains of conidia. 

Family 3. Mortierellacea [Mortierellaceae] Schroter op. cit. 130. Family Chaeto- 
cladiaceae Schroter op. cit. 131. Mucorina whose sporangiophores or conidiophores 
are branched, the sporangia (without columellae) or conidia solitary and terminal 
on the branches. Thamnidium, Chaetocladium, Mortierclla, Haplosporangium. 

Family 4. Zoopagacea [Zoopagaceae] Drechsler in Mycologia 27: 37 (1935). Mu- 
corina parasitic in amoebas or nematodes, producing conidia. The hosts of Zoopaga- 
cea inhabit the soil and are infected by contact with hyphae or conidia. From the point 
of contact, a hypha grows into the host and gives rise to a mycelium; this is in some 
examples reduced to a single coiled cell. The host being killed, the parasite sends 
out hyphae which may produce conidia, usually in chains, or else may conjugate and 
produce zygotes. Endocochlus, Cochlonema, Bdellospora, Zoopage, Acaulopage, 

Family 5. Endogonacea [Endogonaceae] Paoletti in Saccardo Sylloge Fungorum 
8: 905 (1889). Endogonei Fries. Mucorina saprophytic in soil or wood, producing 
macroscopic subterranean fruits. The fruits may reach a diameter of 2 cm. Within 
them, the tips of hyphae are cut off by crosswalls, and develop either into sporangia 
without columellae or into gametes. 

Order 2. Entomophthorinea [Entomophthorineae] (Engler) Campbell Univ. 
Textb. Bot 161 (1902). 
Suborder Entomophthorineae Engler in Engler and Prantl Nat. Pflanzen- 
fam. I Teil, Abt. 1: iv (1897). 
Order Entomophthorales Smith Crypt. Bot. 1: 408 (1938). 
Zygomycetes, mostly parasitic, producing explosively discharged conidia [Masso- 
spora, while clearly belonging to the group, is an exception to the stated character). 
These organisms, although of the general nature of ordinary Inophyta, exhibit 
cytological characters markedly distinguishing the two families from the generality 
of Inophyta and from each other. The position here given to them is the customary 
one; it is doubtful that it is natural. 

Family 1. Entomophthoracea [Entomophthoraceac] Berlese and de Toni in Sac- 
cardo Sylloge 7: 280 (1888). Most species are parasitic in the bodies of insects, 
whose tissues they replace. The hyphae become divided by crosswalls, and the multi- 
nucleate cells thus produced tend to round up and become separate. A well-nourished 
cell may send forth a hypha which reaches the outer air and whose tip is cut off and 
discharged in the direction of the light. Martin (1925) and Couch (1939) described 

Phylum Inophyta [ 125 

the mechanism of discharge. The conidiophore ends in a columella projecting into 
the base of the conidium. The columella develops a double wall. Increasing pressure 
within the conidium causes a sudden eversion of the wall on the side of the conidium, 
and this movement throws the conidium forth to a distance of perhaps 1 mm. Coni- 
dia which come down on unfavorable substrata may form and discharge secondary 

Adjacent cells may conjugate, the thick-walled zygote forming either in one of 
them or as an outgrowth from one of them. Many examples produce thick-walled 
resting spores without conjugation. 

Olive (1906) described the nuclei and the process of mitosis in Empusa. The 
resting nuclei are fairly large, 7-9 [I in diameter. In the course of division, two stain- 
resistant granules are seen, with strands of chromatin radiating from them. These 
move apart, while the nucleus becomes dumb-bell shaped. The nuclear membrane 
remains intact and division is completed by its constriction. As Olive remarked, the 
process is much as in Euglena. 

Entomophthora, Empusa, and Massospora attack insects; the first produces zygotes, 
while the other two produce asexual resting spores; Massospora does not discharge 
the conidia violently. Conidiobolus and Delacroixia are saprophytic. Completoria 
attacks the prothallia of ferns. Ancylistes, a parasite in the green alga Closterium, 
was formerly included among chytrids or Oomycetes. Berdan (1938) showed that it 
belongs here; it produces conidia and zygotes quite of the character of the present 
group, and does not produce zoospores. 

Family 2. Basidiobolacea [Basidiobolaceae] Engler and Gilg Syllab. ed. 9 u. 10: 
45 (1924). Basidiobolus ranarum Eidam (1886) occurs in the intestinal contents of 
frogs and toads as uninucleate cells, solitary or in brief filaments, walled with cellu- 
lose. In manure the filaments develop into a scant branching mycelium. The proto- 
plasm gathers in the ends of erect hyphae which are cut off as conidia and discharged. 
Conjugation occurs between adjacent cells of a filament. It is preceded by a single 
nuclear division in each gamete (Fairchild, 1897). In this process, the nuclear mem- 
brane disappears and the numerous minute chromosomes are found in a blunt-ended 
spindle without centrosomes. Each gamete form a papilla; one of the two nuclei 
enters the papilla, whose contents, after being cut oft by a wall, die and disappear. 
The gametes and their nuclei unite and the zygote secretes a thick wall. 

Class 2. ASCOMYCETES (Sachs ex Bennett and Thistleton-Dyer) 


Order .4jco5porgag Cohn in Hedwigia 11: 17 (1872). 

Ascomyceten Sachs Lehrb. Bot. ed. 4: 249 (1874). 

AscoMYCETES Bennett and Thistleton-Dyer in Sachs Textb. Bot. English ed. 847 

Class AscoMYCETES Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 1 : 32 

Class Ascosporeae Bessey in Univ. Nebraska Studies 7: 295 (1907). 

Class Ascomycetae Schaffner in Ohio Naturalist 9: 449 (1909). 

Inophyta which produce, as a feature of the sexual cycle, sporangia called asci, in 
which the spores, called ascospores, typically eight in number, are delimited by the 
manner of cell division called free cell formation, i.e., in such fashion as to exclude 
a part of the cytoplasm. 

126] The Classification of Lower Organisms 

The hyphae of Ascomycetes are septate and the cells most often uninucleate. 

Most Ascomycetes produce, beside the ascospores, conidia of one type or another. 
A mycelium may produce a mass of densely woven hyphae with conidia on the sur- 
face; such a mass is called an acervulus or sporodochium. Either a mycelium or an 
acervulus or sporodochium may send up spore-bearing columns called coremia. 

Many Ascomycetes produce, either directly from the mycelium or from special 
structures consisting of interwoven hyphae, globular or flask-shaped structures which 
produce conidia internally and release them through a pore. These structures are 
called pycnidia, and the spores pycniospores. In many examples, the pycniospores 
are capable of functioning as sperms; so far as this is true, the pycnidia may alter- 
natively be called spermagonia, and the pycniospores spermatia. 

Hyphae woven into a mass may go into a resting condition, becoming thick-walled, 
hard, and usually dark in color. The resulting structure is a sclerotium. If a structure 
of the general nature of an acervulus, sporodochium, or sclerotium gives rise either 
to pycnidia or to fruits bearing asci, it is called a stroma. 

As to asci and ascospores, Dangeard (1893, 1894, 1907) reached definitely the 
conclusion that they are essentially sexual products. There had been earlier observa- 
tions, beginning with de Bary, 1863, that there are meetings, coilings together, and 
fusions of hyphae as a preliminary to the production of asci. Many ascomycetes are 
of two mating types; this was first discovered of Glomerella, by Edgerton (1914). As 
Dodge (1939) remarks, the mating types are not sexes; in forms producing recogniz- 
able male and female reproductive structures, each mating type may produce both. 

In Ascomycetes which may be regarded as primitive, difTerentiated male and fe- 
male cells are produced. The male cell or antheridium is ordinarily terminal on a 
hypha. The female cell (constituting, together with other differentiated cells of the 
same hypha, if any are present, the ascogonium) may be terminal; more often it bears 
an elongate cell, or a chain of cells, called the trichogyne, and having the function 
of reaching the antheridium. In some Ascomycetes, antheridia are produced, but syn- 
gamy does not take place; the egg is binucleate or multinucleate, and the nuclei 
within it take the part of gamete nuclei in further development. There are others in 
which no antheridia are produced. Hansen and Snyder (1943) found, in Hypornyces 
Solani var. Cucurbitae, that "any part of the living thallus, ascospores, conidia or 
bits of the mycelium could act as the male fertilizing agent." There are forms in 
which fusions take place between undifferentiated hyphal cells; and yet others in 
which it appears that the paired nuclei involved in sexual processes arise by divisions 
of a single nucleus originally present in a spore. 

In some Ascomycetes, syngamy is followed immediately by karyogamy, and the 
zygote develops directly into a single ascus. In the overwhelming majority of the 
group, asci are produced indirectly, and there is no fusion of nuclei until this takes 
place. The zygote sends out hyphae called ascogenous hyphae, recognizably different 
from the vegetative ones. The cells of the ascogenous hyphae arc binucleate; or, 
arising from a multinucleate zygote, become binucleate by the establishment of 
crosswalls. The two nuclei of each cell divide concurrently and the cell walls are so 
placed that each cell receives nuclei of different origin. This effect is achieved in the 
final cell division before ascus formation by a peculiar process called crozier forma- 
tion. The terminal cell of the ascogenous hypha becomes bent to the form of a hook; 
the nuclei divide concurrently, and cell walls appear between the daughter nuclei of 
each pair; the middle cell of the row of three thus produced remains binucleate and 
becomes an ascus. The uninucleate terminal and basal cells lie side by side, and may 

Phylum Inophyta [127 

fuse to form a binucleate cell which may become an additional ascus, or else may 
grow forth and give rise to more asci than one. 

The stage consisting of cells with two nuclei of different origin is called the 
dikaryophase. It is characteristic of Ascomycetes ,and also of Basdiomycetes: among 
Inophyta, it is a normal and familiar thing. To a concept of cytology founded on 
studies overlooking the Inophyta, it would appear an extreme anomaly, almost an 
impossibility. It has the appearance of a rather awkward device for making cells 
genetically and physiologically diploid while the nuclei remain haploid. In most 
Ascomycetes it is a brief stage, but there are some, as Taphrina, whose mycelium 
consists prevalently of binucleate cells. 

The detailed behavior of nuclei in the ascus was first described by Harper (1895, 
1897, 1900) from studies of Peziza, Sphacrotheca, Erysiphe, and Pyronema. The two 
nuclei in the primordium of the ascus unite into one. The fusion nucleus divides 
three, each time in much the same manner. A centrosome with astral rays is 
present at the nuclear membrane, apparently outside. It divides, and a spindle forms, 
inside the intact nuclear membrane, between the daughter centrosomes. The chromo- 
somes appear and divide. As they move toward the poles of the spindle, the nuclear 
membrane collapses or dissolves, leaving the spindle free in the cytoplasm. The 
mass of chromatin at each pole of the spindle shreds out into a nuclear network, 
duly surrounded by a nuclear membrane and usually containing a nucleolus. 

Haploid chromosome numbers of Ascomycetes (all of which have been observed in 
the ascus) include the following: 

Ascoidea rubescens, fide Walker (1935) 2 

Eremascus alhus, fide De Lamater et al. (1953) 6 

G/om^r(?//<z, fide Lucas (1946) 4 

HypornycesSolaniv:xr.Cucurbitae,fi.dtY{.\r?,ch.{\9'^9) .... 4 

Lachnea scutellata,^dt^ro\\'n {\9\\) 5 

Neurospora crassa, fide McClintock (1945) 7 

Peziza do miciliana,^dt ?)c\\n\iz {\921) 8 

Phyllactinia corylea, fide Colson (1938) 10 

Pyronema confiuens\?Lr. igiieum, ^dtV>ro\vn {\9\b) 5 

Taphrina deformans, fide Martin (1940) 4 

According to Harper, when the third division in the ascus is complete, each of 
the eight nuclei produced by it thrusts forths its centrosome upon a beak. The astral 
rays of the centrosomes become recurved in the cytoplasm about the nucleus, and 
grow and multiply until they are converted into a smooth membrane, outside of 
which a wall is deposited. Most observers have not seen so much detail. Brown (1911) 
and Dodge (1937) describe the cell membrane of the ascus, apparently under the 
influence of the centrosome of each nucleus, as cutting into the cytoplasm in an ellip- 
soid pattern. In Taphrina (Martin, 1940), the cytoplasm of the spores is delimited 
simply by accumulation about the nuclei. By whatever process the ascospores are cut 
out. some of the cytoplasm of the ascus is excluded and left without nuclei. Harper 
(1899) proposed to limit the older term free cell formation to processes which have 
this effect; he observed that the occurrence of such processes distinguishes asci from 
the sporangia of Oomycetes and Zygomycetes, in which spores are cut out by cleavage. 
Harper believed that a fusion of nuclei follows immediately the fusion of gametes; 
that the karyogamy observed in the ascus is a uniting of diploid nuclei, producing 
tetraploid nuclei; and that the characteristic three nuclear divisions in the ascus are 
necessary for reduction of the chromosome number from tetraploid to haploid. These 

128] The Classification of Lower Organisms 

hypotheses, long accepted as possible, were disproved by genetic studies by Betts and 
Meyer (1939) and Keitt and Langford (1941). In the asci of many species, the 
spores lie in a single series in which their order is determined by the divisions which 
produce their nuclei. By refined technique, the spores from a single ascus may be 
identified, separated, and cultivated. It is then observed that the mycelia grown from 
the first four spores may differ in some particular character from those grown from 
the second four spores; those from the first pair of spores may differ from those from 
the second; but those from two members of any of the pairs, first, second, third or 
fourth, are always alike. These observations mean that the first two divisions in the 
ascus constitute the meiotic process, the third being mitotic. Lucas (1946) obtained 
cytological evidence refined enough to confirm this conclusion. 

Asci are almost always produced in fruits, which may be called ascocarps. The 
ascocarp aside from the asci arises usually from vegetative hyphae; in the Ascomy- 
cetes regarded as primitive, it does not begin to develop until after fertilization, but 
in the higher ones it may develop in advance of fertilization and become the seat 
of this process. 

There are several types of ascocarps, among which three are most familiar. A 
small ascocarp completely enclosing the asci is a cleistothecium. Cleistothecia were 
formerly included under the term perithecium; that term will better be limited 
to small fruits which are globular or vase-like, opening through a single pore, the 
ostiole, and differing from the pycnidia already described in producing ascospores 
instead of conidia. A fruit in which the asci form a broad layer which is typically 
fully exposed at maturity, the whole being ordinarily of the form of a disk or cup, 
larger than a cleistothecium or perithecium, is an apothecium. 

Asci produced in perithecia or apothecia usually discharge the ascospores vio- 
lently. The mechanism of discharge is apparently simply turgidity. Some asci show 
no visible adaptations for the discharge of spores; others have lids (opercula) 
whose position determines the direction of discharge. Certain large apothecia can 
throw the spores to a distance of 10-20 cm.; the discharge is so governed by tempera- 
ture and humidity as to occur in gently moving rather than in still air. By blowing 
across these apothecia one can make them throw out a visible cloud of spores. 
Heald and Walton (1914) reviewed many older observations of violent discharge by 
perithecia, the oldest by Pringsheim on Sphaeria Scirpi, 1858. Rankin, 1913, found 
that each ascus in turn breaks loose, comes up to the ostiole, projects through it, 
throws out its spores, and collapses to make room for another. Weimer (1920) found 
that the perithecia of Pleurage curvicolla bend toward the light and throw the spores 
to a maximum distance of 45 cm., which is apparently the record. 

There is a widely entertained hypothesis that the Ascomycetes evolved from the 
red algae. It appears to have developed from a piece of classification by Sachs 
(1874), who proposed a class Carposporeen, to consist of the red algae, certain higher 
green algae, and the Ascomycetes and Basidiomycetes. A number of resemblances 
support it. Both red algae and Ascomycetes include many parasites; both lack 
flagellate cells; both have differentiated gametes, the egg bearing a trichogyne; in 
both, fertilization leads to further development before spores are produced. In 
addition to these genuine resemblances, an imaginary one was influential, namely 
the double fertilization ascribed to the red algae by Schmitz and to the Ascomycetes 
by Harper. Numerous as these resemblances are, they are not now believed to indicate 
relationship. Atkinson (1915) formulated the counter-argument. The Ascomycetes 
resemble the Mucorina in nutrition, in producing no flagellate cells, and in multi- 

Phylum Inophyta [ 129 

nucleate gametes. The germination of the zygote of the Mucorina, by the production 
of a hypha bearing a sporangium, resembles the production of ascogenous hyphae by 
the zygotes of Ascomycetes. Two principal changes would convert Mucorina into 
Ascomycetes: the zygote should cease to be a resting spore, and cell division within 
the sporangium should be by free cell formation. This could happen if the centro- 
somes of the ultimate nuclei of the sporangia were in control of cleavage, and if 
these nuclei were so far separated that considerable areas of cell membrane would 
lie beyond the influence of the centrosomes, with the effect that the cell membrane, 
furrowing in to delimit a spore around each nucleus, would leave some of the cyto- 
plasm outside of all of the spores. The organisms listed below as the first order of 
Ascomycetes, Endomycetalea, are but poorly known, yet seem genuinely to represent 
the transition from Mucorina to typical Ascomycetes. 

It is not yet possible to formulate a system of orders of Ascomycetes with the 
expectation that it will not be found to require much amendment^. The following 
will serve tentatively; excellent contemporary authority makes several orders each 
of the ones listed fourth, fifth, and seventh. 
l.Ascus developed directly from the zygote 
(or apomictically from an unfertilized cell); 

not producing fruits Order 1. Endomycetalea. 

l.The zygote giving rise to filaments of cells 
with more than one nucleus, these producing 
the asci. 

2. Producing fruits. 

3. The fruits cleistothecia. 

4. Asci scattered in the fruits; 
mostly saprophytes with 

branched conidiophores Order 2. Mucedines. 

4. Asci in one cluster, or solitary, 
in the fruits; mostly parasites 

with unbranched conidiophores Order 3. Perisporiacea. 

3. The fruits, originally closed, open- 
ing by irregular pores or regular or 

irregular clefts Order 4. Phacidialea. 

3. The fruits apothecia Order 5. Cupulata. 

3. The fruits perithecia. 

4. Producing a normal mycelium Order 7. Sclerocarpa. 

4. Parasitic on insects, the mycel- 
ium reduced Order 8. Laboulbenialea. 

2. Not producing fruits, the asci arising di- 
rectly from the mycelium Order 6. Exoascalea. 

Order 1. Endomycetalea [Endomycetales] Gaumann Vergl. Morph. Pilze 135 

Subclass Hemiasci Engler Syllab. 26 (1892). 

ILuttrell (1951) has presented a complete reorganization of the class. He sets apart 
as a major subordinate group Bitunicatae five orders in which the ripe ascus exudes 
3 vesicle and discharges the spores from this. 

130 ] The Classification of Lower Organisms 

Subclass Hemiasci or Hemiasceae, with suborder (Unterreihe) Hemiascineae, 
and suborder Protoascineae of subclass Euasci, Engler in Engler and Prantl 
Nat. Pflanzelfam. I Teil, Abt. 1 : iv (1897), the names not based on those of 
Order Protoascineae Campbell Univ. Textb. Bot. 165 (1902). 
Order Hemiascalcs Engler Syllab. ed. 3: 28 (1903). 

Ascomycetes whose asci develop directly from the zygotes. Two families may be 

Family 1. Endomycetacea [Endomycetaceae] Schroter in Engler and Prantl Nat. 
Pflanzenfam. I Teil, Abt. 1: 154 (1894). Family Ascoideaceae Schroter op. cit. 145. 
Mostly saprophytes, the uninucleate or multinucleate cells of the filaments tending 
to round up, become separate, and function as conidia; the zygotes, produced by 
syngamy of scarcely differentiated cells, enlarging and becoming asci of 4, 8, or 
many spores cut out by free cell formation. Dipodascus, Eremascus, Endomyces, 
Ascoidea. The asci of the last are apparently produced asexually (Walker, 1935). 

The genus Protomyces requires mention. It is a parasite on higher plants, producing 
walled resting spores which germinate by producing a sporangium of many spores. 
It is chytrid-like, but its spores are non-motile. Its proper place in classification has 
for a long time been a puzzle. 

Family 2. Saccharomycetacea [Saccharomycetaceae] (Rees) Schroter op. cit. 153. 
CXa^?, znd iarm\y Saccharomycetes y<! inter m Rabenhorst Kryptog.-Fl. Deutschland 1, 
Abt. 1: (1879). Unicellular, reproducing by budding, i.e., by production upon 
the cells of outpocketings which are pinched off as additional cells, or by a sexual 
cycle in which endospores are produced, usually by fours. 

These are the organisms which are in English called yeasts. The common bread- 
and beer-yeast called Sac char omyces cerevisiae has a good claim to be considered, 
economically, the most important of all "fungi." Its metabolism, in which dextrose 
is converted to alcohol and carbon dioxide, gives a superficial appearance of sim- 
plicity, and has attracted much study, contributing much to an understanding of the 
genuine intricacy of energesis. 

In addition to agents of fermentation, this family includes pathogens causing 
chronic infections of animals. These have been treated as a genus Torula, Torulopsis, 
Blastodenna, or Cryptococcus. They have not been observed to produce endospores. 

Order 2. Mucedines Fries Syst. Myc. 3: 380 (1832). 

Order Gyjnnoascaceae Winter in Rabenhorst Kryptog-Fl. Deutschland 1, Abt. 

2: 3 (1887). 
Suborder Plectascineae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, 

Abt. 1: V (1897). 
Order Plectascineae Campbell Univ. Textb. Bot. 169 (1902). 
Order Aspergilliales Bessey in Univ. Nebraska Studies 7: 304 (1907). 
Order Gymnascales Clemens Gen. Fung. 93 (1909). 
Order Plcctascales Gaumann Vergl. Morph. Pilze 164 (1926). 
Ascomycetes producing cleistothecia in which the asci are scattered; mostly 
saprophytic and producing branched conidiophores. 

The name Mucedines means molds. Under this name Fries listed twelve genera, 
with Aspergillus Link and Penicillium Link first. The former is the evident standard 
genus of the order. Both genera are very common and numerous in species. They arc 
readily recognized under the microscope by the forms of their clusters of conidia. 

Phylum Inophyta [131 

The conidiophore of Aspergillus ends in a globular swelling from which spring many 
radiating rows of conidia, with the efTect that the entire mass, yellow, brown, black, 
pink, or red in color, is globular. Penicillium has a branching conidiophore bearing 
rows of conidia in a broom-like mass. The masses are usually blue or green, and are 
familiar on cheese, jam, bread, cardboard, oranges, or almost any organic material. 
Particular species of Penicilliuni are involved in the making of genuine Camem- 
bert and Roquefort cheeses. The genus has become best known for the production by 
P. notatum of the drug penicillin. In 1929, Dr. Alexander Fleming of London noticed 
that a mycelium of this species, growing as a contaminant on a plate of bacteria, 
interfered with the growth of the latter. This observation led to the discovery of a 
substance clinically useful against actinomycetes, spheres, and Gram positive rods, 
but not against Gram negative rods. Production was for several years very scant, and 
the drug expensive accordingly; in the early 1940's, as a war measure, the United 
States financed large scale production along with the appropriate scientific study 
(Elder, 1944; Committee on Medical Research, Washington, and the Medical Re- 
search Council, London, 1945). Several forms of penicillin have been recognized; 
they differ in the radicle R in the formula CgHnOiSNiR. The structural formula is 
believed to be as follows (Editorial Board of the Monograph on the Chemistry of 
Penicillin, 1947): 

RCONH — CH — CH — S — C (CH3)2 

I I I 


The sexual reproduction of Aspergillus and Penicillium involves the syngamy 
of differentiated cells. The zygote sends out ascogenous hyphae which bud oflF 
scattered asci; the neighboring cells send out hyphae which become woven into a 
minute firm-walled cleistothecium enclosing them. 

Link, who named Aspergillus and Penicillium., gave to the ascocarp-producing 
stage of Aspergillus the name Eurotium. There is a rule of botanical nomenclature 
which allows only a tentative status to names given to the conidium-producing 
stages of inophytes. Thom and his associates (1926, 1945), in presenting a workable 
system of the species of Aspergillus, remarked that "It is better to forget Eurotium 
along with the technicality." 

This order includes a variety of other molds: Gymnoascus, producing only a 
loose weft of hyphae about the asci; Ctenomyces, on feathers, recognized by comb- 
like outgrowths from the loosely woven ascocarps; Monascus, its name a misnomer, 
the minute fruit containing many asci; Onygena, saprophytic on horns and hoofs, 
producing puffball-like fruits as much as 1 cm. high; Elaphomyces, forming a 
mycorrhiza on roots of conifers and producing hypogaeous fruits as large as walnuts. 

Order 3. Perisporiacea [Perisporiaceae] Fries Syst. Myc. 3: 220 (1829). 
Order Perisporia Fries op. cit. 1: xlviii (1832). 
Suborder Perisporiaceae Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, 

Abt. 2: 21 (1887). 
Subsuborder [Underordnung) Perisporiales Engler in Engler and Prantl Nat. 

Pflanzenfam. I Teil, 1: v'(1897). 
Order Perisporiales Bessey in Univ. Nebraska Studies 7: 295 (1907). 
Ascomycetes producing cleistothecia containing a compact cluster of asci or a 
solitary ascus; mostly parasites producing unbranched conidiophores. 


The Classification of Lower Organisms 

Fig. 25. — Ascomycetes: a-e, Dipodascus albidus after Juel (1902), x 1,000; 
a, gametes; b, syngamy; c, development of ascus; d, e, lower and upper parts of a 
mature ascus. f, Erysiphe graminis, haustorium penetrating an epidermal cell of 
a grass and conidiophore bearing a chain of conidia x 500. g-k, Cleistothecia of 
Perisporiacea x 100: g, of Erysiphe sp.; h, of Microsphaera sp.; i, of Podosphaera sp.; 
j, of Uncinula sp.; k, of Phyllactinia sp. 

Phylum Inophyta [ 133 

The more familiar Perisporiacea are those of family Erysiphea [Erysipheae] 
Winter. They are parasites on plants, mostly producing a white mycelium on the 
surface and sending brief haustoria into the epidermal cells. They produce abundant 
conidia in erect unbranched chains; this habit explains the common name of powdery 
mildews. Harper's important studies of the morphology of Ascomycetes were in large 
part made on powdery mildews. The gametes are uninucleate and unite directly, the 
egg bearing no trichogyne; the ascogenous hyphae are brief; each minute black 
globular cleistothecium bears an equatorial whorl of appendages of a form charac- 
teristic of the genus. In Erysiphe and Sphaerotheca iS. pannosa is the common rose 
mildew), the fruits bear unbranched sinuous appendages like vegetative hyphae; the 
fruit of Erysiphe contains several asci, while that of Sphaerotheca contains one. In 
Microsphaera {M. Alni is the powdery mildew of lilac) and Podosphaera, the ap- 
pendages are dichotomously forked near the tip; the fruit of Microsphaera contains 
several asci, that of Podosphaera only one. The appendages of Uncinula are hooked 
at the tip. Those of Phyllactinia are like sharp spikes with bulbous bases. 

Other Perisporiacea, parasitic or saprophytic on plant material, are compara- 
tively poorly known. The fruits may bear appendages of other characters than those 
of the Erysiphea, or none, and may be characteristically clustered or borne in 
stromata. In some examples the fruits have no definite dehiscence mechanism; in 
others they open by deliquescence or by a separation of plates. Some open by a 
single pore, and appear transitional to those of order Sclerocarpa; some open by a 
cleft, or by lobes separated by radiating clefts, and appear transitional to those of 
order Phacidialea. 

Order 4. Phacidialea [Phacidiales] Bessey in Univ. Nebraska Studies 7: 298 
Phacidiacei Fries Syst. Myc. 1: li (1832). 

Order Hysteriaceae and suborders (of order Discomycetes) Phacidiaceae, 

Stictideae, and Tryblidieae Rehm in Rabenhorst Kryptog.-Fl. Deutsch- 

land 1, Abt. 3: 1, 60, 112, 191 (1896); the ordinal name preoccupied by 

family Hysteriaceae Saccardo. 

Suborders Phacidiineae and Hysteriineae Engier in Engler and Prantl Nat. 

Pflanzenfam. I Teil, Abt. 1 : v ( 1897) . 
Orders Graphidiales and Hysteriales Bessey op. cit. 298, 303. 
Order Hemisphaeriales Theissen in Ann. Myc. 11: 468 (1913). 
Order Microthyriales Clements and Shear Gen. Fung. ed. 2: 94 (1931), 
Ascomycetes producing fruits which are not typical cleistothecia, apothecia, or 

This group is here used as a catch-all for three or more distinct groups, which 
appear to form cross-connections among orders Perisporiacea, Cupulata, and Sclero- 
carpa. This appearance suggests the probability that the present group, and the 
usually accepted orders assembled under it, are not natural, but represent parallel 
developments from several sources. The present groups include moderately numerous 
ordinary parasites and saprophytes, together with great numbers of lichen-formers. 
Only the latter are common and familiar in temperate countries. There has been little 
study of the morphology. 

The families which appear tenable are distinguished as follows: 

a. Fruits minute and flattened, usually releasing the spores through one or more 
pores or clefts (Order Hemisphaeriales Theissen, Microthyriales Clements 
and Shear). 

134] The Classification of Lower Organisms 

Family Microthyriacea [Microthyriaceae] Lindau (in Engler and Prantl, 1897). 
Parasitic on plants, surfaces of the fruits marked by radiating ridges. 

Family Micropeltidacea [Micropeltidaceae] Clements and Shear (1931). Family 
Hemisphaeriaceae Theissen (1913), not based on a generic name. Like the fore- 
going, but the surface of the fruit not radiate or radiate only at the margin. 

Family Trichothyriacea [Trichothyriaceae] Theissen and Sydow. Parasitic on 
inophytes, the mycelium a pseudoparenchymatous layer, asci pendant within the 
fruits from the apparent summit. 

b. Fruits elongate, hard, dark, opening by a narrow cleft (suborder Hysteruneae 

Family Hysteriacea [Hysteriaceae] Saccardo Sylloge 2: 721 (1883). Parasitic on 
higher plants or saprophytic. 

Family Graphidiacea [Graphidiaceae] Clements (1909). An enormous group of 
lichens or parasites on lichens, largely tropical and chiefly crustose, the openings 
of the fruits forming dark lines. 

c. Fruits not as above, mostly with a roundish area of asci exposed by the 
irregular or stellate shattering of a superficial layer; if long and narrow, not 
hard and dark (Suborder PHACiDnNEAE Engler). 

Family Phacidiea [Phacidieae] Saccardo Sylloge 8: 705 (1889). Phacidiaceae 
Saccardo (1889). Family Phacidiaceae Lindau (in Engler and Prantl, 1896). The 
dark fruits thin and weak laterally and below. 

Family Tryblidacea [Tryblidaceae] Rehm (in Rabenhorst, 1896). The dark fruits 
hard and thick laterally and below. 

Family Stictea [Sticteae] Saccardo Sylloge 8: 647 (1889). Stictaceae Saccardo 
(1889). Family Stictidaceae Lindau (1896). Fruits light-colored or white. Higgins 
(1914) found that the agents of the shot-hole disease of plums and cherries, which, 
on the basis of non-fruiting stages, have been called Cylindrosporium Pruni, produce 
on fallen leaves ascocarps distinguishable as three species of the genus Coccomyces 
of the present family. 

Order 5. Cupulata [Cupulati] Fries Syst. Myc. 1 : 2 (1821 ). 

Order Mitrati Fries 1. c; order Uterini Fries op. cit. 1 : liii (1832). 

Family Discomycetes Fries Epicrisis 1 (1836). 

Orders Discomycetes and Tuberaceae Winter in Rabenhorst Kryptog.-Fl. 

Deutschland 1, Abt. 2: 3 (1887). 
Suborders Helevellincae, Pczizineae, and Tuherineae Engler in Engler and 

Prantl Nat. Pflanzenfam. I Teil, Abt. 1: v (1897). 
Orders Helevellincae, Pezizineae, and Tuberineae Campbell Univ. Textb. Bot. 

166, 167, 168 (1902). 

Orders Pezizales, Discolichcnes, Helvellales, and Tuberales Bessey in Univ. 

Nebraska Studies 7: 299, 300, 303, 304 (1907). 

This order includes primarily the cup fungi, the inophytes which produce cup- 

or disk-shaped fruits bearing a single hiyer of closely packed asci on the inner or 

upper surface. There has been much study of some of them, notably of Pyronema, by 

Harper, Dangeard, Claussen, and Brown. The disk-shaped flesh-colored apothecia of 

Pyronema, 1-3 mm. in diameter, are found particularly on damp charcoal. The 

mycelium produces difTcrentiatcd multinucleate antheridia and ascogonia, the latter 

bearing one-celled multinucleate trichogynes. After syngamy, or sometimes without 

it, but always to the best of our knowledge without any fusion of nuclei, the ascogonia 

Phylum Inophyta [ 135 

send out branching filaments which become septate in such fashion that the ultimate 
cells are binucleate. These cells form croziers and produce asci. During the develop- 
ment of the ascogenous hyphae, other hyphae, more slender, grow up from the 
vegetative mycelium; these produce a disk of undifferentiated cells below the layer 
of asci, and send up sterile hairs (paraphyses) among them. 

Gaumann (1926) divided the families of this group into two series by the 
presence or absence of a differentiated operculum at the summit of the ascus. The 
names being put into neuter form, and family Tuberacea being added, the lists are 
as follows: 

Inoperculata : Patellariacea, Dermateacea, Bulgariacea, Cyttariacea, Mollisi- 
acea, Helotiacea, Geoglossacea, Tuberacea. 

Operculata: Rhizinacea, Pyronemacea, Ascobolacea, Fezizacea, Helvellacea. 

Along with these, Clements and Shear (1931) list eight families of lichen-formers, 
some of them very numerous. 

Families Pezizacea and Ascobolacea include the ordinary cup fungi. They are 
mostly saprophytes in soil or on manure, and do not usually produce conidia. Peziza 
was listed by Fries first in order Cupulata; it is the evident standard genus of the order. 

Families Dermateacea and Helotiacea include many parasites on plants. One of 
the Helotiacea is Sclerotinia cinerea, the agent of the brown rot of stone fruits. 
As an active parasite it produces conidia of a type which, if the fruits were unknown, 
would place it in the genus Monilia. These spread the disease rapidly. The killed 
fruits fall and the organism lives in them as a saprophyte, replacing their tissues 
with a hard black mass of hyphae, a sclerotium. This survives the winter and in 
spring sends up stalked white apothecia. 

The Helvellacea have been treated as a separate order, but are not sufficiently 
numerous and distinct to justify this treatment. They are saprophytes in soil, pro- 
ducing large stalked apothecia bearing an extensive layer of asci which is everted 
and wrinkled. The most familiar genera are Elvella and Morchella. The fruits are 
edible, indeed delicious; they should be boiled briefly, then creamed and served on 
toast. When found in abundance they should be preserved by drying for use through- 
out the year. 

The Tuberacea, the truffles, also usually treated as a distinct order, produce 
underground fruits which appear to be apothecia distorted and rolled into balls. 
They are associated with particular species of trees on which the mycelia are be- 
lieved to live as mycorhizae (Dangeard, 1894). The asci commonly contain reduced 
numbers of spores. The fruits are prized by gourmands. 

The relationships of the Cupulata are a puzzle. Pyronema could be interpreted 
as representing an evolutionary transition from the order Mucedines to this. Certain 
parasitic cup fungi produce minute apothecia, hard, dark, and nearly closed, sug- 
gesting a transition to order Sclerocarpa. Some species, particularly among the 
parasites and lichen-formers, seem to intergrade with order Phacidialea, and thence 
again both to Mucedines and Sclerocarpa. The operculate asci which mark a part of 
the group occur also in other orders. Thus there is among Ascomycetes an appearance 
of reticulate relationships, such as reputable naturalists of the past supposed to 
exist in many groups. The appearance is of course illusory; sufficient study of other 
groups has made it possible to distinguish the resemblances among them which indi- 
cate relationship from those which are results of parallel evolution. The study of the 
Ascomycetes has not yet been carried this far. 


The Classification of Lower Organisms 

Fig. 26. — Ascomycetes: a-k, Lachnca scutellata after Brown (1911) x 1,000; 
a, b, formation of crozicr; C, karyogamy; d, fusion nucleus; e-i, stages of mciosis; 
j, k, early stages of free cell formation. 1, Apothecia x 2, and m, ascus x 250, of 
Lamprospora leiocarpa. n, Apothecia x 2, and O, ascus x 250, of Aleuria rutilans. 
V, Apothecia of Sclcrotinia cinerea x 2. q, Fruit x 1, and r, ascus x 250, of Mor- 
chella conica. s-x, Taphrina deformans after Martin (1940) x 1,000; s, growth on 
surface of an infected leaf; t, karyogamy; u, mitosis; V, homeotypic anaphase in the 
ascus; w^ development of ascospores; x^ germination. 

Phylum Inophyta [137 

Order 6. Exoascalea [Exoascales] Bessey in Univ. Nebraska Studies 7 : 305 ( 1907) . 
Suborder Protodiscineae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, 

Abt. 1: V (1897), not based on a generic name. 
Order Protodiscineae Campbell Univ. Textb. Bot. 166 (1902). 
Order Agyriales Clements and Shear Gen. Fung. ed. 2: 141 (1931), in part. 

Ascomycetes parasitic on plants, producing no fruits but a broad layer of asci 
directly on the mycelium. 

The leaves of the hosts of these parasites become swollen and distorted; the 
diseases recognized by these symptoms are called curly-leaf diseases. The most 
familiar is the curly-leaf of peaches, caused by Taphrina {Exoascus) deformans. 
Many others are known. The agents of all of these diseases may be regarded as a 
single family Exoascacea [Exoascaceae] Schroter (in Engler and Prantl, 1894), and 
all are commonly treated as a single genus, Taphrina Fries, typified by T. aurea on 
poplar trees; there are differences among them which might well be treated as of 
generic rank. 

Clements and Shear associated the curly-leaf parasites with a collection of sapro- 
phytes producing small and undifferentiated disk-like or indefinite fruits, as Pyro- 
nema, Ascocorticium, and Agyrium; and offended against the principles of nomen- 
clature by re-naming the order Agyriales. It is probable that something of the nature 
of Agyrium may represent the transition from order Cupulata to this one. 

Martin (1940) described the cytology of Taphrina deformans. The mycelium 
grows between the cells of the host, not penetrating them. It is a dikaryophase 
mycelium, the cells binucleate, the nuclei dividing concurrently, cell division occur- 
ring in such fashion as to separate the daughter nuclei of each pair. In preparation for 
reproduction, hyphae of short round cells form a single layer between the epidermis 
and the cuticle of the host. In each cell of these hyphae, the nuclei unite and then 
divide. The division is mitotic, the fusion nuclei and the daughter nuclei having 
each eight chromosomes. The cell divides, by a wall parallel to the surface of the 
leaf, into two. The daughter cell which lies against the tissues of the host dies, 
and its wall becomes empty; the other cell grows and bursts through the cuticle of 
the host and becomes an ascus. Its nucleus divides three times; the first two divisions 
are the meiotic process, and the chromosome number is reduced to four. Cytoplasm 
accumulates around each of the resulting eight nuclei and is presently cut out by a 
membrane and a wall. No centrosome is evident at any stage of the process. The 
spores germinate by sending out buds, as yeasts form buds; sometimes they do this 
before being discharged from the ascus. So far as Martin could determine, the 
binucleate condition of the mycelium is established by division of the nucleus of 
the spore from which it grows. 

Order 7. Sclerocarpa [SclerocarpiJ Persoon Syst. Meth. Fung, xii (1801). 

Order Pyrenomycetes Fries Syst. Myc. 2:312 (1822); order Uterini, suborder 

Pyrenomycetes Fries op. cit. 1: li (1832). 
Family Pyrenomycetes Fries Epicrisis 1 (1836). 
Order Pyrenomycetes, suborders Hypocreaceae, Sphaeriaceae, and Dothideaceae, 

Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 2: 18, 82, 152, 893 

Suborder {Unterreihe) Pyrenomycetineae, sub-suborders [Unterordnungen) 

Hypocreales, Dothideales, and Sphaeriales Engler in Engler and Prantl Nat. 

Pflanzenfam. I Teil, Abt. 1: v, vi (1897). 


The Classification of Lower Organis?ns 

Order Pyrcnomycetales Bessey in Univ. Nebraska Studies 7: 295 (1907). 
Orders Hypocreales, Sphaeriales, and Dothideales Gaumann Vergl. Morph. 
Pilze 222, 253, 284 (1926). 
Ascomycetes producing, from a normal mycelium, perithecia, i. e., small fruits 
of the shape of a small globe or flask opening through a single pore, the ostiole. 
Sphaeria, which Persoon and Fries listed first under the names which they respec- 
tively used, is the evident standard genus; but this genus has become broken up and 
lost in the work of subsequent scholars. 

This order includes very many species and is by the generality of authority 
divided into three. Forms whose perithecia are borne directly on the m.ycelium, 
together with those whose perithecia are borne in or on but distinct from a dark 

Fig. 27. — Mycosphaerella personata after Higgins (1929), x 1,000; a, conidio- 
phorcs and conidia of Ccrcospora type, b, longitudinal section of pycnidium; 
C, primordium of perithccium with ascogonoium bearing a trichogyne; d, e, ascogen- 
ous hyphae; f, crozier formation; g, longitudinal section of mature peridiecium. 

Phylum Iiiophyta [ 139 

stroma, are assigned to order Sphaeriales. Forms with perithecia in or on a brightly 
colored stroma are Hypocreales. Those whose perithecia are cavities with a wall 
indistinguishable from a dark stroma are Dothideales. These groups are not confi- 
dently acceptable as natural: the stromatic Sphaeriales (Wehmeyer, 1926), the 
Hypocreales, and the Dothideales appear each to include more than one line of 
descent from Sphaeriales with solitary perithecia. 

As a general rule, each perithecium develops in consequence of a separate act of 
fertilization, of a differentiated ascogonium, either by an antheridium, a spermatium, 
or otherwise. 

Gaumann recognized fourteen families in the present group or groups. To these 
are to be added a great number of lichen-formers, properly Sphaeriales and Hypocre- 
ales,. but construed as a single family Verrucariacea; and a smaller number, repre- 
senting the Dothideales, and called Mycoporacea. 

Exmples include the following: 

Among Sphaeriales with solitary perithecia, Mycosphaerella is a genus of more 
than one thousand parasites on plants, mostly inconspicuous, causing leaf spots. 
Their conidia are of various types, Septoria, Phleospora, Ramularia, Cercospora. 
Venturia, another numerous genus, includes V. inaequalis, causing apple scab; its 
conidia are of a type called Fusicladium. 

Four species of Neurospora were discovered by Shear and Dodge (1927) as the 
fruiting stages of a red mold on bread called Monilia sitophila. Genetic study of 
this genus particularly by Tatum, Beadle, and their associates (Ryan, Beadle, and 
Tatum, 1943; McClintock, 1945; Beadle and Tatum, 1945; Tatum and Bell, 1946; 
Mitchell and Houlahan, 1946; Tatum, Barratt, Fries, and Bonner, 1950) has yielded 
results of the highest theoretical significance. Normal cultures require no other 
food than minerals, carbohydrate, and a single vitamin, biotin (Butler, Robbins, 
and Dodge, 1941). Either spontaneously or under violent treatment (with x-rays, 
ultra-violet radiations, or mustard gas) the cultures give rise to many mutations, 
behaving as Mendelian recessives, each consisting of the inability to synthesize 
some one vitamin or amino acid. These observations mean that life in its aspect of 
metabolism consists of unit chemical processes, each controlled by a specific enzyme, 
each enzyme being dependent upon a specific area in a specific chromosome. 

Among stromatic Sphaeriales, Glomerella, with conidial stages identified as 
Gloeosporium or Colletotrichum, attacks many plants; G. cingulata causes the bitter 
rot of apples. Valsa, Diatrype, and Diaporthe are numerous in species. Endothia 
parasitica causes the chestnut blight, destructive in the eastern United States. 
Xylaria, Daldinia, and other genera are saprophytic on wood; the former produces 
black fruits, club-shaped or branched; the latter, fruits of the form of black knobs 
which may reach the size of golf balls. 

Among Hypocreales, Nectria cinnabarina is common as a saprophyte on dead 
twigs of poplar. It produces small wart-like red stromata which bear first conidia, 
then perithecia. Claviceps purpurea causes a disease of rye; it produces conidia of 
various types, and converts the grains of rye into sclerotia. These bodies are called 
ergot; they are extremely poisonous, sometimes dangerously so, because they may 
be ground with the grain. They are used in medicine. After lying in the earth through 
the winter, the sclerotia send up fruits of the form of a stalk bearing a knob consisting 
of radiating perithecia. Cordyceps kills subterranean larvae or pupae of insects and 
then sends up a stalk bearing an elongate head of many perithecia. 

140 ] The Classification of Lower Organisms 

The Dothideales include Plowrightia morhosa, the agent of the black knot of 
plums. Diseased twigs become swollen and covered with a black stroma which bears, 
according to the season, conidia of various types or else perithecia. 

Order 8. Laboulbenialea [Laboulbeniales] Engler Syllab. ed. 3: 42 (1903). 
Order Laboulbeniaceae Thaxter, the name (ascribed to Peyritsch) preoccupied 

by family Laboulbeniaceae Berlese in Saccardo Sylloge 8: 909 (1889). 
Suborder Laboulbeniineae Engler in Engler and Prantl Nat. Pflanzenfam. 

I Teil, Abt. 1: vi (1897). 
Class LabouJbeniomycctes Engler Syllab. 1. c. 
Class Laboulbenieae Schaffner in Ohio Naturalist 9: 450 (1909). 

Parasites on insects, the mycelium scant or reduced to a single cell, producing 
antheridia which discharge spermatia into the air and small numbers of perithecia. 

These organisms have the appearance of excep^^ional setae on their hosts, which 
are not usually seriously injured by them. They were first mentioned in a note by 
the entomologist Rouget, 1850; Montagne and Robin, in Robin's book on parasitic 
plants, 1853, gave the first names, Laboulbenia Rougetii and L. Guerinii, the generic 
name honoring the entomologist Laboulbene. Only a few scholars, notably Thaxter 
(1896, 1908, 1924, 1926, 1931) have given much attention to this group; they have 
distinguished well over a thousand species, forming three families and about fifty 

Many Laboulbenialea occur as two forms, male and hermaphrodite. A male indi- 
vidual produces a series of flask-shaped antheridia, each of which discharges into 
the air, one at a time, a series of globular naked sperms. A hermaphrodite individual 
produces first a series of antheridia as described and then one or more perithecia. 
A perithecium consists of a wall, of a definite number of cells produced in definite 
order and pattern, surrounding an egg which bears a trichogyne; the trichogyne 
protrudes from the perithecium and receives the sperms. The zygote gives rise to a 
fascicle of asci which crowd aside and destroy the inner cells of the wall and dis- 
charge the ascospores (usually eight in the ascus, and divided into two cells) through 
the ostiole. 

Those who would link the Ascomycetes with the red algae entertain the hypothesis 
that the Laboulbenialea represent the transition. This hypothesis is surely mistaken. 
The Laboulbenialea are a highly specialized group, not a link between others. They 
appear to have evolved from Sphaeriales with solitary perithecia. 

Class 3. HYPHOMYCETES Fries 

Classes Hyphomycetes and Coniomycctes Fries Syst. Myc. 3: 261, 455 (1832). 

Families Hyphomycetes and Coniomycctes Fries Epicrisis 1 (1836). 

Fungi imperfecti or Deuteromycetes Auctt. 

Inophyta of which the structures involved in sexual reproduction are unknown. 

It has been noted that a particular genus of Ascomycetes may produce conidia 
of more types than one, as Sclcrotinia produces types called Monilia and Botrytis, 
and Glomerella produces types called Gloeosporium. and Colletotrichum. The same 
type may be produced by many genera; the Monilia type recurs in Neurospora, 
which does not belong to the same order as Sclerotinia. Collecting naturalists, and 
plant pathologists in the pursuit of their duties, are constantly encountering conidial 
stages whose assignment to an order of Ascomycetes is impossible. It is an obvious 

Phylum Inophyta [ 141 

practical necessity that a register of these observations be kept. The register is pro- 
vided by the present group, one which is named, defined, and assigned to the category 
of classes, and divided into named orders, families, and genera under which specimens 
may be identified as of species old or new. Class, orders, families, and genera are 
known not to be valid taxonomic groups; many of the ostensible species are known, 
and most of the rest are believed, to be stages of organisms which would in other 
stages have other names. Almost all of them are Ascomycetes; Zygomycetes and 
Basidiomycetes do not usually occur in unidentifiable stages. 

The ascus-bearing stages are constantly being discovered. When this happens, the 
species is re-named in its proper place among Ascomycetes. Theoretically, it loses 
its place in the list of imperfect fungi; practically, it retains it, because the next 
collector or plant pathologst is most likely to try to find it there. 

The system of Hyphomycetes is as follows: 

Order 1. Phomatalea [Phomatales] Clements Gen. Fung. 121 (1909). 
Sphaeropsideae Saccardo Sylloge 8: xvi (1889). 
Order Sphaeropsidales Engler in Engler and Prantl Nat. Pflanzenfam. I Teil 

Abt. 1**: v (1900), not based on a generic name. 
Order Phomales Clements and Shear Gen. Fung. ed. 2: 175 (1931). 

Producing pycnidia. The four families correspond with as many groups of Ascomy- 

Family 1. Phomatacea [Phomataceae] Clements Gen. Fung. 121 (1909). Family 
Sphaerioideae or Sphaerioidaceae Saccardo; but Sphaeria belongs to order Sclero- 
carpa. Family Phomaceae Clements and Shear (1931). Pycnidia hard and black as 
in Sphaeriales and Dothideales. Phoma, Ascochyta, Diplodia, Septoria, each of 
many species. 

Family 2. Zythiacea [Zythiaceae] Clements Gen. Fung. 128 (1909). Family 
Nectrioideae or Nectrioidaceae Saccardo; but Nectria belongs to order Sclerocarpa. 
Pycnidia in brightly colored stromata as of Hysteriales. 

Family 3. Leptostromatacea [Leptostromataceae] Saccardo Sylloge 3: 625 (1884). 
Pycnidia in shield-like stromata, like the fruits of Microthyriacea. 

Family 4. Discellacea [Discellaceae] Clements and Shear Gen. Fung. ed. 2: 192 
(1931). Family Excipulaceae Saccardo; but Excipula is a cup fungus. Pycnidia wide 
open like the fruits of Phacidiea. 

Order 2. Melanconialea [Melanconiales] Engler in Engler and Prantl Nat. Pflan- 
zenfam. I Teil, Abt. 1**: v (1900). 

The conidia borne on a stroma but not in pycnidia. 

Family Melanconiacea [Melanconiaceae] (Saccardo, without category) Lindau 
in Engler and Prantl op. cit. 398, the single very numerous family: Gloeosporium; 
Coryneum, C. Beijerinckii, the shot-hole of almonds; Pestallozia. 

Order 3. Nematothecia [Nematothecii] Persoon Synops. Meth. Fung, xix (1801). 
Orders Dematiei, Sepedoniei, Tubercularini, and Stilhosporei Fries Syst. Myc. Order 
Hyphomycetes (Fries) Auctt. Order Moniliales Clements Gen. Fung. 138 (1909). 
Conidia directly on the mycelium, or none. 

Family 1. Tuberculariea [Tubercularieae] Saccardo Sylloge 4: 635 (1886). 
Tuberculariaceae Saccardo (1889). Family Tuherculariaceae Lindau (1900). 
Scarcely distinct from Melanconiacea, the conidia on a mass of interwoven hyphae 

142 ] The Classification of Lower Organisms 

less compact than a stroma. Fusarium, an enormous number of species producing as 
conidia crescent-shaped rows of cells. Snyder and Hansen (1941, 1945) find that 
the fruiting stages are species of Hypomyces, Nectria, Gibberella, or Calonectria, all 

Family 2. Stilbellacea [Stilbellaceae] Bessey Morph. and Tax. Fungi 584 (1950). 
Family .Siz/^e'a^ Saccardo Sylloge 4: 563 ( 1886). .S^z/foacfflP Saccardo ( 1889). Family 
Stilbaceae Lindau (1900); Bessey observed that the type of the genus Stilbum does 
not belong to this family. Mostly molds producing coremia. 

Family 3. Dematiea [Dematieae] Saccardo Sylloge 4: 235 (1886). Dematiaceae 
Saccardo (1889). Family Dematiaceae Lindau (1900). Dark-colored parasites, as 
Helminthosporium, Cladosporium, and Cercospora, or molds, as Alternaria. 

Family 4. Moniliacea [Moniliaceae] Clements Gen. Fung. 138 (1909). Mucedineae 
Persoon, family Mucedineae or Mucedinaceae Saccardo, not based on a generic 
name. White or brightly colored parasites or molds, as Oidium, with colorless spores 
in chains, Monilia, Botrytis, etc. The parasites on animals which have been referred 
to Monilia are currently called Candida. 

Family (?') 5. Sterile mycelia. Many mycorhizae must be left here. Rhizoctonia, 
dark net-like masses of hyphae occurring as parasites or saprophytes. Trichophyton, 
parasitic on the skins of man and animals, causing ringworm, athlete's foot, etc. 

Class 4. BASSDIOMYCETES (Sachs ex Bennett and Thistleton-Dyer) 


Order Basidiosporeae and subordinate group Basidiomycetae Cohn in Hedwigia 
11: 17 (1872). 

Basidiomyceten Sachs Lehrb. Bot. ed. 4: 249 (1874). 

Basidiomycetes Bennett and Thistleton-Dyer in Sachs Textb. Bot. English ed. 
847 (1875). 

Class Basidiomycetes Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 
1: 72 (1884). 

Classes Teliosporeae and Basidiosporeae Bessey in Univ. Nebraska Studies 7 : 305, 
306 (1907). 

Classes Teliosporeae and Basidiomycetae Schaffner in Ohio Naturalist 9 : 450 

Inophyta which produce, as a feature of the sexual cycle, conidiophores called 
basidia, each producing typically four conidia called basidiospores. 

Germinating basidiospores give rise to mycelia of cells with solitary haploid 
nuclei. Syngamy occurs among cells of these mycelia, usually simply by contact of 
vmdifTerontiated cells; the rusts produce differentiated sperms in spermagonia re- 
sembling the pycnidia of Ascomycetes. In some species any haploid hypha may 
conjugate with any; in some there are two mating types, and in some four. Raper 
(1953) has studied the interesting genetics of the mating types. 

The cell produced by .syngamy remains undifferentiated, but gives rise, by con- 
current division of its nuclei, to a dikaryote mycelium. The nuclei are minute, and 
mitosis has rarely been seen. The nuclear divisions are often followed by a peculiar 
manner of cell division, comparable to the crozier formation of Ascomycetes, and 
producing structures called clamp connections. 

Either the original haploid mycelium or the dikaryophase may produce conidia 
without nuclear change. Such reproduction is familiar among the rusts, rather un- 
familiar among other Basidiomycetes. 

Phylum Inophyta [ 143 

Only the dikaryophase produces the specialized conidiophores called basidia, 
which are regularly the seat of karyogamy and meiosis. There is a considerable 
variety of types of basidia. Van Tieghem (1893) originated the terminology ap- 
plicable to these; Martin (1938) has attempted to refine it, and Linder (1940) to 
simplify it. 

Frequently, the seat of meiosis is a thick-walled resting spore or an otherwise 
difTerentiated cell called a probasidium, upon which the proper basidium develops, 
after meiosis, as an outgrowth. A basidium arising in this fashion is commonly 
elongate and divided into four cells each of which produces a basidiospore. Such a 
hypha-like basidium may be called a promycelium or a phragmobasidium; the latter 
term is applicable also to an elongate four-celled basidium which does not arise 
from a probasidium. In a few Basidiomycetes, the basidium is divided into four cells 
by longitudinal walls; such basidia are called cruciate basidia. In the familiar 
Basidiomycetes the basidium does not become divided by walls and is called a holo- 
basidium or autobasidium. Gaumann ( 1926) distinguished two types of holobasidia: 
the stichobasidium, in which the spindles of the dividing nuclei lie at various levels 
and in various directions, and which frequently produces more than four nuclei; 
and the chiastobasidium, in which the spindles lie transversely near the summit, 
and which regularly produces just four nuclei. Dodge, translating Gaumann (1928), 
denies much importance to this distinction. 

The meiotic divisions have repeatedly been studied. Apparent centrosomes have 
been seen at the poles of the spindles (Lewis, 1906; Lander, 1933), but not by most 
microtechnical methods (Savile, 1939; Ritchie, 1941). The chromosomes gather as 
usual at the middle of the spindle and divide. The nuclear membrane becomes in- 
distinct, but the nuclear sap remains distinct from the cytoplasm nearly until the 
completion of division; it then disappears, leaving the groups of daughter chromo- 
somes connected by a spindle of the appearance of a dark streak in the cytoplasm. 
Ob:;erved haploid chromosome numbers include the following: 

Coleosporium, fideMoxczM (1914) 2 

Coleosporium Vernoniae, fide Olive (1949) 8 

Coj&rmuj, fide Yokes (1931) 4 

Eocronartium, fide Fitzpatrick (1918) 4 

&;frffa, fide Whelden (1935) 4 

Gymnosporangium, fide Stevens (1930) 2 

Melampsora, fide Savile (1939) 4 

Myxomycidium flavum, fide Martin (1938) 8 

Puccinia, fide Savile (1939) 4 

Transchelia, fide Savile (1939) 4 

Russula, fide Ritchie (1941) 4 

Scleroderma, fide Lander (1933) 2 

f/romycg'^, fide Savile (1939) 4 

Savile suggests that some at least of the reports of a chromosome number of 2 
may have resulted from misinterpreted observations of one pair of choromosomes 
behind another. 

Normally, only the two meiotic divisions, producing four nuclei, occur in the 
basidium; exceptionally, there are further, mitotic, divisions, resulting in more than 
four spores on the basidium. The basidiospores are usually borne on slender stalks 
called sterigmata. Sterigmata and spores are formed by evagination of the wall of 
the basidium; the nuclei migrate through the sterigmata into the spores. 


The Classification of Lower Organisms 

Fig. 28. — Basidiomygetes : a. Two germinating basidiosporcs of Agariciis campcs- 
tris produce mycelia which anastomose freely, the cells becoming plurinucleate, 
after Hein ( 1930) , x 500. b, c. Young and older basidia of Cystobasidium sebaceum, 
after Martin (1939). d-g, Eocronartium muscicola after P'itzpatrick (1918); d, fus- 
ion nucleus; e, homeotypic division in the basidium; f, four-celled basidium; g, pro- 
duction of basidiospore. h^ i^ i, Basidia of Ustilago Heujlcri, U. Hurdei, and Tille- 

(Continued bottom p. 145) 

Phylum Inophyta [ 145 

Most basidia discharge the spores actively, to a distance of a fraction of a milli- 
meter. Buller (1929) observed that just before a spore is cast off a minute droplet 
of liquid appears at the summit of the sterigma. This occurs in precisely the same 
fashion in mushrooms, rusts, certain smuts, and the yeast-like organism Sporoholo- 
myces. Buller inferred that the force which discharges the spore is surface tension 
in the droplet. The fruits of Basidiomycetes are evidently adapted to the feebleness 
of the mechanism by which the spores are discharged. If the fruits are cup-like, 
they open laterally or downward. The basidia of mushrooms stand horizontally on 
gills which are commonly less than one millimeter apart, allowing the spores to 
fall from between them without touching them. 

The groups of Ascomycetes and Basidiomycetes are evidently related. Morels and 
mushrooms, truffles and puffballs, taste alike. The technical scholar will be con- 
vinced that the groups are related by the occurrence in both of a dikaryophase 
stage, a character too strongly in contrast with those of the generality of organisms 
to be a probable product of parallel evolution. Gaumann quotes an old opinion of 
Vuillemin (1893), "qu'une baside est un asque dont chaque cellule-fille avant de 
passer a I'etat de spore, fait saillie au dehors et se transforme en une sorte de conidie 
pour mieux s'adapter au transport par la vent." In dealing with the Zygomycetes, 
Gaumann emphasized the apparent evolution of conidia from endospores by evagina- 
tion of the walls of the sporangia. Largely, as it seems, by Gaumann's influence, 
Vuillemin's hypothesis has become generally accepted. 

Gaumann was disposed to derive the Basidiomycetes from something like Asco- 
cortkiiim, and began his account of several of the groups of Basidiomycetes with 
forms having scant flat fruits, or having basidia which spring directly from the 
substratum or host. Linder (1940) suggested a derivation from Cupulata or Sclero- 
carpa having operculate asci. He took note that many such asci open by producing a 
vescicle, bounded by the stretched inner wall of the ascus, into which the asco- 
spores pass. This led to the conclusion that the Basidiomycetes producing probasidia 
are the lowest, and to this extent his reasoning appears cogent. He went on to identify 
the rusts as the lowest Basidiomycetes, which seems far-fetched, the rusts being 
distinctly a specialized group. 

The generally accepted groups of Basidiomycetes are those which were set forth 
by Engler (1897, 1900), as follows: 

Subclass HEMiBAsron, having basidia bearing indefinite numbers of spores; the 

Subclass EuBAsron, the basidia bearing definite numbers of spores. 

Order {Reihe) Protobasidigmycetes, the basidia divided into cells. 

Suborder {Unterreihe or Ordnung) ^uricularhneae, the basidia divided 
by transverse walls. 

Sub-suborder [Unter ordnung) Uredinales, the rusts. 

tia Tritici, after Sartoris ( 1924) . k, 1, Basidia of Patouillardina cinerea after Martin 
(1935). m, Basidium of Sebacina sublilacina after Martin (1934). n, Basidium 
of Protodontia Uda after Martin ( 1932). o, p, younger and older basidia of Tulas- 
nella phaerospora, after Martin (1939). q-t, Development of the basidium of 
Guepinia Spathularia, after Bodman (1938). u-x, Russula emetica after Ritchie 
(1941); binucleate primordium of basidium, fusion nucleus, homeotypic division, 
development of basidiospores. y, z, Basidia of Lycogalopsis Solmsii after Martin 
(1939). X 1,000 except as noted. 

146 ] The Classification of Lower Organisms 

Sub-suborder Auriculariales. 
Suborder Tremellineae, the basidia divided by longitudinal walls. 
(At this point should appear Reihe Autobasidiomycetes, to include eight 
Unterreihen of ordinary Basidiomycetes. The name Autobasidiomycetes 
does not appear in the table of contents, the text, or the index of the 
Natilrlichen Pflanzenfamilien; it was published in Engler's Syllabus, 1892). 
Rearranging these groups according to current opinion, and suppressing the sub- 
sidiary categories, one arrives at the following system of orders: 
1. Producing probasidia or transversely divided 
basidia, usually both. 

2. Probasidia, if formed, terminal on the 

3. Mostly saprophytic and producing 

gelatinous fruits Order 1. Protobasidiomycetes, 

3. Parasitic, mostly not producing 

fruits; the rusts Order 2. Hypodermia. 

2. Probasidia produced by rounding up and 
deposition of thiclc walls by the gener- 
ality of the cells of the mycelium Order 3. Ustilaginea. 

1. Without probasidia, the basidia divided lon- 
gitudinally Order 4. Tremellina. 

1. Without probasidia, the basidia undivided. 
2. Fruits gelatinous, basidia producing only 

two spores on stout sterigmata Order 5. Dacryomygetalea. 

2. Not as above. 

3. Basidia in a layer which forms with- 
out protection or becomes exposed. . .Order 6. Fungi. 
3. Basidia formed in closed fruits 
which do not open to expose them 
as a single layer Order 7. Dermatocarpa. 

Order 1. Protobasidiomycetes Engler in Engler and Prantl Nat. Pflanzenfam. I 
Teil, Abt 1**: iii (1900). 
Suborder Auriculariineae and sub-suborder Auriculariales Engler 1. c. 
Order Auricularineac Campbell Univ. Textb. Bot. 175 (1902). 
Order Auriculariales Bcssey in Univ. Nebraska Studies 7: 309 (1907). 
Basidiomycetes mostly producing probasidia, th.^ basidia divided by transverse 
walls, mostly saprophytic and producing gelatinous fruits. 

This order includes the family Auriculariacea [Auriculariaceae] Lindau in Engler 
and Prantl Nat. Pflanzenfam. I Teil, Abt. 1**: 83 (1900), from which two or three 
others have been segregated; about fifteen genera and about 125 species. 

Martin (1943) has discussed the name of the genus Auricularia and of its type 
species. The organism in question is surely the Jew's ear, Tretnella Auricula L.; the 
genus Auricularia Bulliard 1795 can have nothing else as a type. The right name 
of the species is Auricularia Auricula (L. ) Underwood 1902. It is a saprophyte on 
logs and sticks, producing flattened brown gelatinous fruits a few centimeters in 
diameter, vaguely resembling human ears. There are no probasidia. Hyphae growing 
toward the surfaces of the fruits produce a palisade of elongate basidia. Each basi- 
dium becomes divided by transverse walls into four cells, and each of these sends out 

Phylum Inophyta [ 147 

to the surface an elongate sterigma which bears a curved basidiospore. The organism 
produces also conidia, either from the mycelium, the fruits, or directly from the 

A series of unfamiliar other genera, Platygloca, Cystobasidium, Septobasidium, 
etc., have been studied notably by Martin (1934, 1937, 1939, 1942). Jola and 
Eocronartium are parasites on mosses. All of these genera produce probasidia, from 
which four-celled phragmobasidia arise, as a layer near the surfaces of the fruits. 
Most of them produce also conidia. 

Order 2. Hypodermia [Hypodermii] Fries Syst. Myc. 3: 460 (1832). 

Uredinees Brongniart in Bory de Saint Vincent Diet. Class. Hist. Nat. 16: 471 

Order Uredineae Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 1: 
74 (1884). 

Sub-suborder Uredinales Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, 
Abt. 1**: iii (1900). 

Order Uredinales Bessey in Univ. Nebraska Studies 7: 306 (1907). 

Order Pucciniales Clements and Shear Gen. Fung. ed. 2: 147 (1931). 
The rusts: parasitic Basidiomycetes, the haploid and dikaryote mycelia usually 
attacking different hosts; the dikaryote mycelium producing probasidia, these not 
usually compacted into fruits, usually heavily walled and serving as resting spores, 
becoming or giving rise to four-celled phragmobasidia. 

The typical reproductive structure of the haploid stage is the aecium, a cup-shaped 
structure which releases spores called aeciospores; this stage is accordingly called the 
aecial stage, and its host the aecial host. In addition to aecia, this stage usually pro- 
duces pycnidia or spermagonia. The typical reproductive structures of the dikaryote 
mycelium are clusters (telia) of spores called teliospores or teleutospores; this stage, 
then, is the telial stage, and its host the telial host. The telial stage usually produces, 
beside the teliospores, others called uredospores. The teliospore, or rather (since the 
teliospore commonly consists of two or more cells) each cell of the teliospore, is a 
probasidium, producing a promycelium which bears four basidiospores. These state- 
ments mean that a normal rust produces spores of five kinds. Rusts producing differ- 
ent kinds of spores were formerly supposed to be different genera; such were the 
Aecidium, Uredo, and Puccinia of Persoon, who, however, remarked of Uredo line- 
aris, "vereor, ne junior plantula Pucciniae graminis modo sit." De Bary first proved 
that Aecidium Berberis is yet another stage of Puccinia graminis. 

The dikaryophase is initiated, of course, by syngamy among cells of the aecial 
stage. In Phragmidium violaceum, Blackman observed this to take place between 
different cells of the same hypha. Christman (1905) and Moreau (1914), studying 
other species of Phragmidium, observed fusion to take place between tips of different 
hyphae. Craigie, 1927, showed that Puccinia graminis occurs in two mating types, 
and that the fertilizing elements are pycniospores or spermatia. De Bary (1884) had 
suggested that this is the truth; his suggestion waited some forty years to be confirmed. 
Allen (1930) has described much of the detail. The pycniospores are carried out of 
the pycnidium in exuding fluid, and are carried by insects; they make protoplasmic 
connection with paraphyses growing from pycnidia of the opposte mating type. The 
binucleate uredospores arise from a dikaryote mycelium, but the cup-shaped wall of 
the aecium is produced by the haploid mycelium. 

148 ] The Classification of Lower Organisms 

The first-formed reproductive structures of the dikaryote mycelium on the telial 
host are usually uredospores, which remain binucleate and have the function of 
spreading the infection of the telial host. 

Teliospores may be compacted into palisade-like masses which break through the 
epidermis of the host; the masses may be gelatinous and yellow, like fruits of 
Auriculariacea. In other genera, the teliospores are gathered into hard, microscopic- 
ally stout columns, and in yet others they break through the epidermis in masses not 
compacted, each teliospore on a separate stalk. The teliospores of Phragynidium are 
chains of several probasidia; those of the many species of Pucciriia are chains reduced 
to two probasidia; those of Ravcnelia are globular clusters of probasidia. Almost 
always, the teliospores are thick-walled; outside of the tropics, they have the function 
of overwintering. Each probasidium contains two nuclei. These unite as a preliminary 
to germination: this was first observed by Sappin-Troufi^y (in Dangeard and Sappin- 
Troufi'y, 1893). Thereafter the probasidium gives rise to the four-celled promycelium. 

The life cycle thus described is not perfectly stable. Aeciospores, uredospores, and 
young teliospores are alike dikaryote, and are genetically identical. Spores of the 
structure and behavior of any of these types may be produced by processes which 
normally lead to another. Thus in Puccinia Malvaccaruvi, the hollyhock rust, syng- 
amy leads directly to the production of teliospores on the host of the haploid mycel- 
ium; spermagonia, aecia, and uredosori are not produced. 

Four families of rusts may be recognized (various authorities make fewer or more). 
There are about five thousand species. 

Family 1. Melampsoracea [Melampsoraceae] Dietel in Engler and Prantl Nat. 
Pflanzenfam. I Teil, Abt. 1**: 38 (1900). Teliospores forming a single compact 
layer and germinating by producing promycelia. The aecial stages are mostly on 
conifers. Some have telial stages on ferns, and FauU (1929) regards these as most 
primitive; others attack a variety of flowering plants. 

Family 2. Coleosporiacea [Coleosporiaceae] Auctt. The teliospores themselves 
becoming basidia by transverse division. In some examples, as Gallowaya, they are 

Family 3. Cronartiacea [Cronartiaceae] Auctt. The teliospores compacted into 
columns. Cronartium, with aecial stages on pines; C. ribicola, the important white 
pine blister rust, its telial stage on gooseberries and currants. 

Family 4. Uredinacea [Uredinaceae] Cohn in Hedwigia 11: 17 (1872). Family 
Pucciniaceae Dietel op. cit. 48. The bulk of the rusts, producing teliospores on indi- 
vidual stalks. Hemileia vastatrix, the coffee rust; Phragmidium spp., autoecious (at- 
tacking a single host) on Rosaceae; Gymnosporangium, the aecial stage on junipers, 
the telial (with no uredospores) on plants of the apple tribe; Puccinia, a great num- 
ber of species. The races which attack barberry and grasses are all called Puccinia 
graminis; but there are morphologically distinguishable strains on wheat, rye, oats, 
timothy, Agrostis, and blue grass. Leading an active sexual life and capable of muta- 
tion, these strains are subdivisible into large numbers of races distinguished by capa- 
city to attack different races of hosts. Given a specimen of rust on wheat, one deter- 
mines by trial upon seedlings of ten varieties of wheat to which of 189 numbered races 
it belongs. The races occur characteristically in different wheat-growing areas. If one 
breeds wheat for resistance to rust, there is good probability of success against the 
races occurring locally; but some other race is likely to move into the area (Stakman, 

Phylum Inophyla [ 149 

Order 3. Ustilaginea [Ustilagineae] (Tulasne and Tulasne) Winter in Rabenhorst 
Kryptog.-Fl. Deutschland 1, Abt. 1: 73 (1884). 
Ustilagineae Tulasne and Tulasne in Ann. Sci. Nat. Bot. ser. 3, 7: 73 (1847). 
Subclass Hemibasidii Engler Syllab. 26 (1892). 
Order Ustilaginales Bessey in Univ. Nebraska Studies 7: 306 (1907). 

The smuts: parasitic Basidiomycetes completing their development on a single 
host, the dikaryophase mycelium breaking up into thick-walled black spores, these 
functioning as probasidia, the basidia usually bearing more than four basidiospores. 

In the apparently more primitive smuts, the promycelia are four-celled phragmo- 
basidia. The haploid nuclei divide before passing into the basidiospores, with the 
effect that each cell of the promycelium buds off a series of basidiospores. In other 
examples the promycelia do not become divided by walls, but are of the character of 
holobasidia. The basidiospores of some species are capable of budding like yeasts. 
In some species, they are capable of syngamy with each other, and in some they send 
out hyphae which bear conidia of characteristic form. In many species, syngamy has 
not been observed, but is beheved to take place between vegetative hyphae. Hybridi- 
zation, and mutation, particularly in the capacity to attack particular races of hosts, 
take place freely in smuts, which are accordingly well fitted to cope with the efforts 
of plant breeders. 

The smuts are believed to be somewhat degenerate descendants of the rusts. 

There are two families, about thirty genera, about six hundred species. 

Family 1. Ustilaginacea [Ustilaginaceae] Cohn in Hedwigia 11: 17 (1872). The 
basidia divided by transverse walls. Ustilago, on grasses and other plants. 

Family 2. Tilletiacea [Tilletiaceae] Dietel in Engler and Prantl Nat. Pflanzenfam. 
I Teil, Abt. 1** : 15 ( 1900) . The basidia not divided by walls. Tilletia, on grains, etc. 
Tuburcinia, Doassansia, the resting spores produced in globular masses. 

Order 4. Tremellina [Tremellinae] Fries Syst. Myc. 1: 2 (1821); 2: 207 (1822). 
Order Tremellinei Fries Hymen. Eur, 1 (1874). 
Order Tremellineae Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 1 : 

74 (1884). 
Suborder Tremellineae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, 

Abt. 1**: iii (1900). 
Order Tremellales Bessey in Univ. Nebraska Studies 7: 309 (1907). 
Order Tulasnellales Gaumann Vergl. Morph. Pilze 487 (1926). 
Saprophytic Basidiomycetes producing gelatinous fruits bearing a layer of basidia 
which typically become divided into four cells by longitudinal walls. Each cell pro- 
duces a long stout sterigma which reaches the surface of the fruit and bears a spore. 
The mycelia, the young fruits, or the basidiospores may bear conidia. 

The number of species is perhaps one hundred. Nearly all belong to family Tre- 
mellacea [Tremellaceae] Cohn in Hedwigia 11: 17 (1872). Martin (1935, 1937, 
1939) has given much study to this group. It is clearly related to the Protobasidi- 
omycetes; Patouillardina, having basidia divided by oblique walls, is clearly transi- 
tional. Tremella, Sebacina, Tremellodendron, Hyaloria. 

Tulasnella differs from the generality of Tremellina in producing holobasidia of 
a peculiar type, with bulbous sterigmata (Lindau interpreted the sterigmata as basi- 
diospores borne without sterigmata and not released, but producing conidia; it may 
be that this interpretation is more sound than the obvious one). It is supposed that 
the holobasidia of this genus are derived from the cruciate basidia of proper Tremel- 

150 ] The Classification of Lower Organisms 

lina by a line of descent separate from those which have produced the holobasidia of 
other groups. By leaving Tulasnella in order Tremellina, we spare ourselves the recog- 
nition of one more insignificant order. 

Order 5. Dacryomycetalea [Dacryomycetales] Gaumann Vergl. Morph. Pilze 490 
Suborder Dacryomycetineae Engler in Engler and PrantI Nat. Pflanzenfam. 
ITeil, Abt. 1**: iv (1900). 

Saprophytic Basidiomycetes producing small gelatinous fruits bearing holobasidia 
in which two of the nuclei produced by meiosis undergo degeneration, while two 
pass into the basidiospores by way of stout sterigmata which give the basidium the 
form of a Y. Conidia are produced either from the mycelium, from the young fruits, 
or from the basidiospores. 

There is a single family Dacryomycetacea [Dacryomycetaceae] Hennings in Engler 
and PrantI Nat. Pflanzenfam. I Teil, Abt. 1**: 96 (1900). Dacryomyces, Dacryomi- 
tra, Guepinia. Bodman ( 1938) observed the details of the cytological processes in the 

This insignificant order, like Tulasnella and the two great orders next to be con- 
sidered, is evidently derived from Protobasidiomycetes, through Tremellina, by loss 
of septa in the basidia; the peculiarities of its basidia suggest an independent origin. 

Order 6. Fungi L. Sp. PI. 1171 (1753). 

Order Hynienothecii Persoon Syst. Meth. Fung, xvi (1801). 

Class Hymenomycetes and orders Pilcati and Clavati Fries Syst. Myc. 1: 1, 2 

Yzmily Hymenomycetes Fries Espicrisis 1 (1836). 
Family Agaricaceae Cohn in Hedwigia 11: 17 (1872). 
Order Hymenomycetes Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 

1: 74 (1884). 
Suborders Exobasidiineae and Hymenomycetineae Engler in Engler and PrantI 

Nat. Pflanzenfam. I Teil, Abt.'l**: iv (1900). 
Orders Hymenomycetales and Exohasidiales Bessey in Univ. Nebraska Studies 

7: 307, 308 (1907). 
Order A^aricalcs Clements Gen. Fung. 102 (1909). 

Orders Cantharellales, Polyporales, and Agaricales Gaumann Vergl. Morph. 
Pilze495, 503, 519 (1926). 
Basidiomycetes producing holobasidia in a layer which is or becomes exposed to 
the air, usually on fruits which are woody, leathery, or fleshy, rather than waxy or 

The layer of basidia is called the hymenium. In the lowest members of the group, 
the hymenium is formed directly on the mycelium, on the surface of the host or 
substratum; in higher examples, it is formed on the surface of more or less compli- 
cated fruits; in the highest, it is formed in closed fruits which open to expose it. The 
area of the hymenium, and the number of basidia it can bear, is increased when it is 
not smooth, but thrown into teeth, ridges, plates, or other projections. Families have 
been distinguished chiefly on the basis of the form of the hymenium. The system is not 
reliably entirely natural; Overholts (1929) pointed out various microscopic details 
which promise to contribute to a more natural system. Among these are cystidia, 
swollen cells imbedded in the hymenium and projecting from it; in some examples at 

Phylum Inophyta [151 

least, they are sterile basidia and serve to hold apart the ridges bearing the hymenium. 
Other microscopic features are setae, similar to cystidia but hard, dark, and pointed; 
slender hairs called paraphyses; latex ducts; and crystalline inclusions. 

There are some fifteen thousand species. The following famiHes are for the most 
part the conventionally accepted ones. 

Family 1. Exobasidiacea [Exobasidiaceae] Hennings in Engler and Prantl Nat. 
Pflanzenfam. I Teil, Abt. 1**: 103 (1900). The basidia directly on the mycelium. A 
sniall group, mostly parasitic on plants. Exohasidium. 

Family 2. Thelephoracea [Thelephoraceae] (Saccardo) Hennings (1900). Order 
Thelephorei Fries Hymen. Eur. 1 ( 1874) . FamHy Thelephorei Winter ( 1884) . Thele- 
phoraceae Saccardo Sylloge 8: xiii (1889). Fruits of various form, gelatinous, fleshy 
or leathery, the hymenium covering the surface generally except where it faces up- 
ward. Corticium, saprophytic, the fruit a mere appressed layer; Stereum, leathery 
shelf-like extensions from decaying sticks and logs: these genera seem to lead into 
Lnmily Polyporacea. Cora, a tropical variant of Stereum, is the only lichen-forming 
basidiomycete. Thelephora, Craterellus, the fruits club-, funnel-, or cup-like. 

Family 3. Clavariacea [Clavariaceae] (Saccardo) Hennings (1900). Order 
Clavariei Fries (1874). Family Clavariei Winter (1884). Clavariaceae Saccardo 
(1889). Fruits fleshy, club-like or branched; stag-horn fungi. Clavaria, generally 

Family 4. Hydnacea [Hydnaceae] (Saccardo) Hennings (1900). Order Hydnei 
Fries (1874). Family //yi/n^i Winter (1884). Hydnaceae Saccardo (1889). Hymen- 
ium on the surface of downward-pointing teeth. Fruits assigned to the genus Hydnum 
may be massive or variously branched or mushroom-shaped, leathery or fleshy; the 
fleshy examples are edible. Fruits of Irpex are little leathery brackets projecting from 
sticks and logs, distinguished from Stereum or Polystictus by the masses of fine teeth 
projecting below. 

Family 5. Polyporacea [Polyporaceae] (Saccardo) Hennings (1900). Order Poly- 
poreiYries (1874). Family Polyporei Winter (1884). Polyporaceae Saccardo (1889). 
The hymenium lining vertical tubes open below. These are mostly woody or leathery 
shelf fungi, mostly saprophytic on wood, numerous and varied in detail. Cooke ( 1940) 
recognized forty-six genera in North America. Polyporus, Fames, Polystictus. In Dac- 
dalea, the pores are not cylinders but slits; this genus leads into Lenzites, in which the 
hymenium is borne on radiating plates, and which is conventionally stationed in 
Agaricacea. Boletus has stout fleshy mushroom-shaped fruits, yellow to brown, turn- 
ing green when bruised. These fruits are unattractive, but some species are eaten; 
others are supposed to be poisonous. 

Family 6. Agaricacea [Agaricaceae] Cohn in Hedwigia 11: 17 (1872). Order 
Agaricini Fries (1874). Family Agaricini Winter (1884). The hymenium on vertical 
plates, radiating from a center, called gills. 

These are the Fungi whose fruits are called mushrooms or toadstools. The fruits 
are mostly mushroom-shaped, sometimes shelf-like; the texture is usually fleshy, vary- 
ing to leathery on the one hand, and on the other to deliquescent, i.e., becoming 
converted after maturity into black fluid. There has been much study of the develop- 
ment of the fruits (Levine (1922) and Hein (1930) give extensive bibliographies). 
This occurs in any of several different fashions, leading to recognizable differences in 
the mature structure. For the identification of agarics, many mushroom books are 
available. Any interested person, noting the details of structure which result from 
the different courses of development, together with the color of the spores (of one 

152 ] The Classification of Lower Organisms 

of five classes, white, pink to red, light brown to rust color, dark brown or purple, or 
black), will find identification reasonably easy. Popular interest in agarics is con- 
cerned, of course, with the edible and poisonous. Many amateur mycophagists need to 
be convinced that there is no single test for poisonous agarics except the final one. 
One who encounters an unfamiliar species may chew and eat a small scrap of it; if 
it is tasty and without bad after-effects, one may collect and eat the same species 
when one again recognizes it by its technical characters. At the present point, it is 
expedient to mention only a few examples. 

Deliquescent agarics with black spores are called inky caps and constitute the genus 
Coprinus. All are edible; they should be fried in butter and served on toast. 

Fruits of Agaricus campestris, the field mushroom, are rather large, white or gray 
on top, the stalk marked by a ring but no cup, the gills pink when young, dark brown 
to nearly black when mature. Anything of this character is safely edible. 

Fruits of Pleurotus have an excentric or lateral stalk, or none, being shelf- or 
bracket-like, fleshy, with white spores. All species are edible. The most familiar is 
the oyster mushroom, P. ostreatus, producing large white to gray fruits on dead 
trees, commonly on poplars. 

Fruits of Amanita are marked by cup and ring, and bear white spores. Some species 
are known to be edible; others, as the fly agaric, A. muscaria, recognized by a red cap 
flecked with white, are extremely poisonous. 

Family 7. Podaxacea [Podaxaceae] Fischer in Engler and Prantl Nat. Pflanzenfam. 
I Teil, Abt. 1**: 332 (1900). Gyrophragmium produces fruits much like those of 
Agaricus, but coming up only to ground level, and drying and shattering irregularly 
instead of opening like mushrooms. The gills are quite evident in immature fruits. 
Podaxon is similar, but does not form definite gills. These organisms are convention- 
ally stationed in the next order, but their obvious natural position is next to 

Order 7. Dermatocarpa [Dermatocarpi] Persoon Syst. Meth. Fung, xiii (1801). 
Order Lytothccii Persoon op. cit. xv. 
Class Gasteromycetes and orders Angiogastres and Trichospermi Fries Syst. Myc. 

2: 275, 276 (1822). 
Family Gasteromycetes Fries Epicrisis 1 (1836) 
Order Gasteromycetes Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 

Suborders Phallineae, Hymenogastrineae, Lycoperdineae, Nidulariineae, and 
Plectobasidiineae Engler in Engler and Prantl Nat. Pflanznfam. I Teil, Abt. 
1**: iv (1900). 
Orders Phallineae, Lycoperdineae, and Nidularineae Campbell Univ. Textb. 

Bot. 186, 187, 188 (1902). 
Orders Hymenogastrales, Phallales, Lycoperdales, Nidulariales, and Sclcroder- 

matales Bessey in Univ. Nebraska Studies 7: 306-307 (1907). 
Orders Plectobasidiales and Gasteromycetes Gaumann Vergl. Morph. Pilze 537, 
Basidiomycetes producing holobasidia enclosed in fruits, not forming a continuous 
layer or not exposed as such, not discharging the spores directly into the air, sterig- 
mata more or less suppressed. 

Distinguished by negative characters, this order may be suspected of being artificial; 
but Engler's attempt to correct this produced orders which were small and numerous 

I'lixUnu Innfihytd 

[ 153 

Fig. 29. — Fruits of Agaricacea: upper left, Coprinus atramcntarius; upper right, 
Galera tenera; below, Agaricus campestris. Photographs by the late Dr. J. J. McCabe, 
by courtesy of the Department of Botany, University of California. 

Phylum Inophyta [ 155 

to an unsatisfactory degree, and to some of which the suspicion of artificiality con- 
tinued to attach. 

Dodge, translating Gaumann (1928), took account of the course of development 
of the fruits in rearranging those families whose fruits are characteristically pro- 
duced underground. The roll of families which appear tenable is as follows. 

A. Fruits typically formed underground. 

Family 1. Rhizopogonacea [Rhizopogonaceae] Dodge in Gaumann Comp. Morph. 
Fungi 469 (1928). 

Family 2. Sclerodermea [Sclerodermei] Winter in Rabenhorst Kryptog.-Fl. 
Deutschland 1, Abt. 1: 865 (1884). Family Sclerodermataceae Fischer in Engler 
and Prantl Nat. Pflanzenfam. I Teil, Abt. 1**: 334 (1900). 

Family 3. Hydnangiacea [Hydnangiaceae] Dodge in Gaumann op. cit. 485. 

Family 4. Hymenogastrea [Hymenogastrei] Winter in Rabenhorst op. cit. 865. 
Family Hymenogastraccae de Toni in Saccardo Sylloge 7: 154 (1888). 

Family 5. Hysterangiacea [Hysterangiaceae] Fischer in Engler and Prantl op. cit. 

B. Fruits appearing on the surface of the ground. 

Family 6. Lycoperdacea [Lycoperdaceae] Cohn in Hedwigia 11: 17 ( 1872) . These 
are the common puffballs, Lycoperdon, Bovista, Calvatia, Lycogalopsis, etc. The con- 
tents of the more or less globular fruits become disorganized, leaving a mass of spores 
m.ixed with fibers (modified hyphae constituting a capillitium), enclosed in one or 
more continuous layers of tissue (peridia) which open usually through one stellate 
pore at the summit. Geaster has a double peridium. The outer peridium becomes 
split by meridional clefts from the apex nearly to the base, and the lobes curl back 
in damp weather, exposing the inner peridium with its terminal pore. The appearance 
of the fruit in the damp condition explains the common name, earth star, and the 
scentific name of the same meaning. 

Family 7. Tulostomea [Tulostomei] Winter in Rabenhorst op. cit. 866. Family 
Tulostomataceae Fischer in Engler and Prantl op. cit. 342. Tulostoma produces at 
ground level puffball-like fruits which are found to stand upon buried stalks some 
centimeters long. The basidia bear the spores scattered along the sides instead of in 
a crown at the summit. This is probably a minor deviation from the condition in 
ordinary puffballs, and not a token of independent origin. 

Family 8. Nidulariea [Nidulariei] Winter in Rabenhorst 1. c. Family Nidulariaceae 
de Toni in Saccardo Sylloge 7: 28 (1888). The bird's nest fungi, Nidularia, Cyathus, 
etc., with small fruits growing on sticks or earth, the outer peridium opening and 
exposing several peridioles. 

Family 9. Sphaerobolacea [Sphaerobolaceae] Fischer in Engler and Prantl op. 
cit. 346. Sphaerobolus, a saprophyte on wood, produces minute puffball-like fruits 
which discharge mechanically a globular mass of spores. 

Family 10. Clathracea [Clathraceae] Fischer in Engler and Prantl op. cit. 280. 
Closely related and transitional to the following family. 

Family 11. Phalloidea [Phalloidei] Winter in Rabenhorst 1. c. Family Phallaceae 
Fischer in Engler and Prantl op. cit. 289. The stinkhorns. Phallus, Dictyophora, 
Mutinus, etc. These organisms produce highly specialized fruits. A fruit is first seen 
as a white globe, as large as a marble or a golf-ball, at ground level. It has a leathery 
peridium containing certain structures imbedded in gelatinous matter: there is a 
firm thimble-shaped structure upon whose surface the basidia develop; below or 
within this there is a body of the form of a hollow cylinder of spongy structure. When 

156 ] The Classification of Lower Organisms 

the spores are ripe, the spongy body grows, so to speak, by unfolding, and becomes, 
it may be within an hour, a stalk as much as 15 cm. tall. This happens usually during 
the night or at dawn, and is not commonly observed. The growing stalk carries the 
basidium-bearing structure into the air, bursting the peridium, which remains as a 
cup about the base, and exposing the spores in a mass of jelly which is of an odor 
repulsive to man but attractive to carrion-seeking insects. The latter are used as 
agents of dissemination. 

Chapter X 

Phylum 6. PROTOPLASTA Haeckel 

Stdmme Protoplasta and Myxomycetes Haeckel Gen. Morph. 2: xxiv, xxvi 

Subphylum Plasmodroma Doflein Protozoen 13 (1901), in part. 
Subphylum Rhizoflagellata Grasse Traite Zool. 1, fasc. 1: 133 (1952), not order 

Rhizoflagellata Kent (1880). 
Further names for the myxomycetes as a phylum are cited below under class 

Organisms without photosynthetic pigments, mostly with flagellate stages, the 
flagella simple or acroneme, not paired and equal nor solitary and posterior; com- 
monly occurring also in amoeboid stages. By Haeckel's original publication, the type 
or standard is Amoeba, i.e., Amiba diffluens. 

Amoeboid organisms are those whose protoplasts lack walls or shells, or are only 
incompletely covered by them, and which thrust forth temporary bodies of proto- 
plasm, called pseudopodia, functional in motion and in predatory nutrition. Pseudo- 
podia are of several types. If massive and blunt they are lobopodia. If fine and 
straight, not anastomosing and usually not branching, they are filopodia; or, if they 
contain inner filaments, axopodia. If fine, branching, and anastomosing, they are 

The characters of the pseudopodia distinguish the accepted primary groups of 
amoeboid organisms. Variations in this character tend to run parallel to variations 
in the structure and composition of shells and skeletons: to a considerable extent, 
the accepted groups appear natural. This applies to the second, third, and fourth 
among the classes treated below. The phylum, on the other hand, is acknowledgedly 
artificial. Some of its groups appear to have had their origins (presumably more 
origins than one) among the chrysomonads; others are of unguessed origin. 

1. Flagellate in the vegetative condition Class 1. Zoomastigoda. 

1. Amoeboid in the vegetative condition. 

2. Producing rhizopodia; with shells, these 

usually calcareous Class 3. Rhizopoda. 

2. Producing filopodia or axopodia; mostly 

with skeletons, these usually siliceous Class 4. Heliozoa. 

2. Producing lobopodia. 

3. Producing flagellate reproductive 

cells; mostly macroscopic, subaerial Class 2. Mycetozoa. 

3. Not as above; without flagellate 

stages Glass 5. Sarkodina. 

Class 1 . ZOOMASTIGODA Calkins 

Subclass Zoomastigina Doflein Lehrb. Prot. ed. 4: 462 (1916). 
Class Zoomastigoda Calkins Biol. Prot. 285 (1926). 
Class Zooflagellata Grasse Traite Zool. 1, fasc. 1: 574 (1952). 
Class Zoomastigophorea Hall Protozoology 170 (1953). 

158] The Classification of Lower Organisms 

Non-pigmented flagellates having acroneme or simple flagella; amoeboid stages, 
if they occur, having lobopodia. The standard is Bodo. Four orders are to be recog- 

1. Flagella one or two Order 1. Rhizoflagellata. 

1. Flagella four to eight (in each neuromotor 
system, if these are more than one). 

2. Axostyles, if present, homologous with 
flagella; parabasal body commonly ab- 
sent Order 2. PoLYMASXiGroA. 

2. Axostyles present, not homologous with 
flagella; parabasal body present, disap- 
pearing during mitosis Order 3. Trichomonadina. 

1. Flagella of indefinite large numbers Order 4. Hypermastigina. 

Order 1. Rhizoflagellata [Rhizo-Flagellata] Kent Man. Inf. 1: 220 (1880). 

Orders Trypanosomata (the mere plural of a generic name) and Flagellato- 

Pantostomata in part Kent op. cit. 218, 229. 
Suborders Monadina in part and Heteromastigoda Biitschli in Bronn Kl. u. Ord. 

Thierreichs 1: 810, 827 (1884). 
Protomastigina Klebs in Zeit. wiss. Zool. 55: 293 (1893). 
Order Protomonadina Blochmann Mikr. Tierwelt ed. 2, 1 : 39 (1895). 
Subclasses Pantostomatineae and Protomastigineae Engler in Engler and Prantl 

Nat. Pflanzenfam. I Teil, Abt. la: iv (1900). 
Orders Pantostomatales and Protomastigales Engler Syllab. ed. 3: 7 (1903). 
Orders Cercomonadinea and Monadidea in part Poche in Arch. Prot. 30: 139, 

140 (1913). 
Orders Pantostomatineae and Protomastigineae Lemmermann in Pascher Siiss- 

wasserfl. Deutschland 1: 30, 52 (1914). 
Order Rhizomastigina Doflein Lehrb. Prot. ed. 4: 704 (1916). 
Orders Pantostomatida and Protomastigida Calkins Biol. Prot. 286, 288 (1926). 
Orders Trypanosomidea Grasse, Bodonidea Hollande, and Proteromonadina 

Grasse in Grasse Traite Zool. 1, fasc. 1: 602, 669, 694 (1952). 

Orders Rhizomastigida and Protomastigida Hall Protozoology 171, 173 (1953). 

Non-pigmented flagellates with one flagellum or two unequal flagella, these 

simple or acroneme; commonly with amoeboid stages, or amoeboid while bearing 

flagella. The type, being the sole genus of Rhizo-Flagellata as originally published, is 

Mastigamoeba, i. e., Chaetoproteus Stein. 

As the synonymy shows, most authorities have made these organisms two orders, 
Pantostomatales (or some such name), amoeboid in the vegetative condition, and 
Protomastigina (or the like), not definitely so. Monas, and the choanoflagellates 
and Amphimonadaceae, usually included in the latter order, have in the present work 
been given places elsewhere. The residue of the Protomastigina are not sharply 
different in character from the original Rhizoflagellata, and are accordingly placed 
in the same order. The resulting group is not a very numerous one. Some examples 
appear to occur naturally as predators in uncontaminatcd waters; the majority have 
been found in foul or contaminated waters, or in feces, and are believed to be 
naturally cntozoic, cither commensal or parasitic. Further examples are parasites in 
blood. A cytological character marking the majority of the goup, but not confined 
to it, is the parabasal body (better, perhaps, the kinetoplast; Kirby, 1944). This is a 

Phylum Protoplasta [ 159 

rather massive extranuclear body regularly present in the cell and distinct both 
from the centrosome and the blepharoplast. In the present group, it divides when 
the nucleus does. Thus this group, although marked chiefly by characters which are 
negative or derived, appears possibly to be natural. 

1. Flagella two. 

2. Cells not notably slender Family 1. CERCOMONADroA. 

2. Cells notably slender Family 2. TRYPANOPLASMroA. 

1. Flagellum one. 

2. Not regularly markedly amoeboid Family 3. Oicomonadacea. 

2. Conspicuously amoeboid Family 4. CHAETOPROXEroA. 

Family 1. Cercomonadida [Cercomonadidae] Kent Man. Inf. 1: 249 (1880). 
Family Bodonina Butschli in Bronn Kl. u. Ord. Thierreichs 1: 827 (1884). Family 
5oc?onflccag Senn in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. la: 133 (1900). 
Family Bodonidae Doflein Protozoen 73 (1901). Family Cercobodonidae Hollande 
1942. Family Proteromonadidae Grasse Traite Zool. 1, fasc. 1: 694 (1952). Non- 
pigmented flagellates, the bodies not notably slender, with two flagella, one directed 
anteriorly, the other trailing. Fischer (1894) found both of the flagella of Bodo to 
be acroneme. 

In Bodo both flagella are free of the body. There are numerous species, in infusions 
or foul or polluted waters, or entozoic in a wide variety of animals, from insects to 
men. Prowazekia, Proteromonas, and Pleuromonas are doubtfully c'istinct. Rhyncho- 
monas, from fresh or foul waters, is distingished by a protoplasmic beak in which 
the anterior flagellum is imbedded. Cercomonas, of like habitats, has the trailing 
flagellum grown fast to the cell membrane; the eel! : exhiljit a considerable capacity 
to send out lobopodia. 

Biflagellate organisms which can lose their flagella and take on the appearance 
of ordinary amoebas have repeatedly been discovered and variously named. So far as 
the pseudopodia are lobopodia and the flagella are unequal, these organisms belong 
in this family; but many accounts fail to establish the equality or inequality of the 
flagella, with the result that the names used in them cannot be applied with confidence. 
This is true of various organisms originally named under Pseudospora, Dimastiga- 
moeba, and Naegleria. The earliest generic name definitely applicaple to organisms 
as described in Cercobodo Senn, 1910. 

Belar (1914, 1916, 1920, 1921), Kuhn (1915), and others have described mitosis 
in various examples of this family; the most detailed account is of Bodo Lacertae in 
Belar's paper of 1921. The flagella spring from a blepharoplast from which a rhizo- 
plast extends into the nucleus. The chromatin is reticulate, not massed in a karyo- 
some, but no centrosome has been recognized in it when it is not dividing. The 
rhizoplast, where it passes through the cytoplasm, is surrounded by stainable Ring- 
korper. The parabasal body, located on the posterior side of the nucleus, is massive 
and often irregular. In division, the blepharoplast divides, each part retaining one 
flagellum and generating an additional one. The rhizoplast appears to begin to split, 
but presently it and the Ringkorper become invisible. Within the intact nuclear mem- 
brane there appears a spindle with evident centrosomes at the poles. The centrosomes 
come presently to the inner surface of the nuclear membrane, while the blepharo- 
plasts move to adjacent positions on the outside. Chromosomes duly assemble at 
the equator of the spindle and undergo division. Division of the nucleus is com- 
pleted by constriction of the nuclear membrane; the parabasal body undergoes 
constriction; the cell divides by constriction lengthwise. The Ringkorper and the 
rhizoplast are apparently regenerated by the blepharoplast. 


The Classification of Lower Organisms 


Fig. 30. — Rhizoflagellata : a, Bodo sp. x 1,000. b, c, Cercomonas longicauda 
as identified by Wenyon (1910) in material from a cholera patient; d, the same as 
identified by Hovasse (1937) in swamp water, e-h, Cryptobia spp.; e-g, cell and 
division stages of a species from the conger eel after Martin (1910); h, a species 
from siphonophores after Keysselitz (1904) x 1,000. i, Phytomonas Donovani after 
Franga (1914). j-p, Trypanosoma Lewisi; j, k, forms from the rat after Minchin 
(1909); 1-p, forms from the flea Ceratophyllus fasciatus after Minchin & Thomp- 
son (1915). q. Division stage of Trypanosoma Brucii after Kiihn & Schuckmann 
(1911). r, Chaetoproteus [Mastigamoeba aspera) after Schulze (1875) x 100. 
X 2,000 except as noted. 

Phylum Protoplasta [161 

AlexeiefF (1924) described fusions of pairs of cells of Bodo edax. 

Family 2. Trypanoplasmida [Trypanoplasmidae] Hartmann and Jollos 1910. Fam- 
ily Cryptobiidae Poche in Arch. Prot. 30: 148 (1913). Family Trypanophidae Hol- 
lande in Grasse Traite Zool. 1, fasc. 1: 680 (1952). Organisms of essentially the 
structure of Cercomonas, but notably slender in adaptation to parasitic life, the 
trailing flagellum forming the margin of an undulating membrane on the body. 
Parasitic in various invertebrates and in the gut and blood of fishes. 

The numerous species may be included in a single genus Cryptobia Leidy [Try- 
panoplasma Laveran and Mesnil; Trypanophis Keysselitz). 

According to Martin's (1910) description of a species from the eel Conger niger, 
both flagella spring from a blepharoplast ("basal granule") at the anterior end. 
As preliminary to division, the blepharoplast and flagella divide, and one blepharo- 
plast migrates to the posterior end of the cell. The nucleus divides by constriction of 
the nuclear membrane. There is a prominent parabasal body ("kinetonucleus") which 
divides by constriction, as does the cell, transversely. 

Belar (1916) described sexual fusions of differentiated individuals of a species 
parasitic in snails. 

Family 3. Oicomonadacea [Oicomonadaceae] Senn in Engler and Prantl Nat. 
Pflanzenfam. I Teil, Abt. la: 118 (1900). Family Trypanosomidae Doflein Proto- 
zoen 55 (1901). Family Trypanosomatidae Grobben 1904. Family Oicomonadidae 
Hartog. Non-pigmented anteriorly uniflagellate organisms, not markedly amoeboid 
while in the flagellate condition. 

Oikomonas includes organisms of the character of the family without particular 
specialization, occurring in contaminated water or soil, and as commensals in the 
intestine of animals. 

The bulk of the family consists of the slender-celled parasites which may be 
celled trypanosomes in the broad sense of the word. From the viewpoint of man, these 
are the most important flagellates, and they have been the most intensely studied. 
Some are known only from the guts of insects; some occur alternatively in insects 
and plants; some in insects and vertebrates; and some in vertebrates and in inverte- 
brates other than insects, as ticks and leeches. The range of parasitization is as 
though the group had evolved as parasites in insects, and had been carried to 
other hosts by the activity of insects and other biting or sucking invertebrates. 

Most trypanosomes occur in varied forms. The forms are designated by words 
which originated as names of genera and remain in use as such. ( 1 ) The leptomonas 
form has an anterior flagellum but no undulating membrane; it resembles a cell of 
Oikomonas but is notably slender. (2) The leishmania form has no flagellum; the cell 
is rounded up and lives attached to, or inside of, cells of the host. (3) In the crithidia 
form, the base of the flagellum is continued as an undulating membrane more or less 
to the middle of the cell. (4) In the trypanosoma form, the base of the flagellum is 
continued as an undulating membrane to the posterior end of the cell. 

The accepted genera are distinguished (artificially, as one may suspect) by stages 
produced and groups of hosts attacked, as follows: 

l.With leptomonas stages in insects and in 
Euphorbiaceae, Ascelepiadaceae, and other 
plants with milky juice Phytomonas. 

1. Confined to invertebrate animals. 

2. Trypanosoma stage known Herpetomonas. 

2. Trypanosoma stage unknown; crithidia 

162 ] The Classification of Lower Organisms 

stage known Crithidia. 

2. Trypanosoma and crithidia stages un- 
known Leptomonas. 

1. Attacking vertebrate animals. 

2. Trypanosoma stage known Trypanosoma. 

2. Trypanosoma stage unknown Leishmania. 

Man has been concerned particularly with Trypanosoma gambiense, the agent of 
African sleeping sickness; T. Cruzi, the cause of Chagas' disease; T. Brucii, T. Evansi, 
T. equinum, and T. equiperdum, which cause in domestic animals the diseases, 
respectively, nagana, surra, mal de caderas, and dourine; Leishmania Donovani and L. 
tropica, causing kala azar and oriental sore; and L. brasiliensis, causing espundia, 
ferida brava, or chicleros' ulcer, usually appearing as a grievous disfigurement of 
the features. 

Schaudinn (1903), having studied a trypanosome occurring in mosquitoes and in 
the owl Athene noctua, described the nucleus as undergoing repeated unequal divi- 
sions. It appeared to him that when a cell is to produce a flagellum, one of the 
minor nuclei produced by unequal division generates it. Prowazek (1903) described 
similar phenomena in a Herpetomonas occurring in flies. These reports led Woodcock 
(1906) to apply to the proper nucleus of trj^panosomes the term trophonucleus, and 
to the large granule near the base of the flagellum the term kinetonucleus. 

There has been much other study of the cytology of trypanosomes (as by Minchin, 
1908, 1909; Robertson, 1909; Woodcock, 1910; Minchin and Woodcock, 1910, 1911; 
Kiihn and Schuikmann, 1911; Minchin and Thomson, 1915; Schuurmans Stekhoven, 
1919). This has not confirmed the foregoing accounts and conclusions, but appears 
to have established the following points. 

The base of the flagellum is slightly swollen and may be construed as a blepharo- 
plast. Separated from the blepharoplast by a distance of one or two microns there 
is a conspicuous parabasal body (the kinetonucleus of Woodcock). Fine strands con- 
necting the blepharoplast, parabasal body, and nucleus, have been observed. Most of 
the stainable material in the resting nucleus is aggregated in a globular karyosome. 
In mitosis, the karyosome breaks up to form a moderate number of chromosomes and 
a central granule, evidently a centrosome, which stains more heavily than the chromo- 
somes. It divides before the chromosomes, the daughter centrosomes remaining con- 
nected by a fine fiber, the centrodesmose. An obscure spindle forms about the centro- 
desmose; thi' chromosomes undergo division within the spindle, and the daughter 
chromosome > assemble about the centrosomes. Mitosis is completed by constriction 
of the nuclear membrane. 

The blepharoplast divides at the same time as the nucleus. The flagellum splits 
to a short distance and one of the branches breaks loose; one daughter blepharoplast 
retains essentially the whole of the original flagellum while the other generates one 
which is almost entirely new. The parabasal body undergoes constriction. The cell 
membrane cuts in in such fashion as to divide the cell longitudinally. The blepharo- 
plast and the parabasal body persist through the non-flagellate leishmania stage. 
Reports that the nucleus may generate these structures, or that one of them may 
generate another, were apparently mistaken. 

Schaudinn described complicated processes by which a trypanosome generates 
differentiated male and female gametes which duly undergo syngamy. His account is 
believed to have resulted from mistaking stages of a sporozoan for those of a trypa- 
nosome. Still, the occurrence of syngamy among trypanosomes is inherently probable. 

Phylum Protoplasta [ 163 

Family 4. Cliaetoproteida [Chaetoproteidae] Poche in Arch. Prot. 30: 172 (1913). 
Family Rhizomastigina Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 810 (1884). 
Family Rhizomastigaceae Senn in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 
la: 113 (1900). Family Mastigamoebidae Kudo Protozoology ed. 3: 263 (1946). 
Amoeboid organisms bearing one anterior flagellum, either permanently or tempor- 
arily. In polluted soil or water, or commensal or pathogenic in animals. 

The oldest genus, Chaetoproteus Stein {Mastigamoeba F. E. Schulze, 1875; Din- 
amoeba Leidy ?) remains poorly known. This organism and Mastigella are described 
as fairly large; Craigia is much smaller. Rhizomastix is doubtfully distinct from 
Craigia. Early names of this family appear to refer to Rhizomastix as the type, but 
the family is much older than the genus, and the names are not valid. 

Order 2. Polymastigida Calkins Biol. Prot. 292 (1926). 

Family Polymastigina Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 842 (1884). 
Order Polymastigina Blochmann Mikr. Tierwelt ed. 2, 1: 47 (1895). 
Subclass Distomatineae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, 

Abt. la: iv (1900). 
Order Distomatinales Engler Syllab. ed. 3: 7 (1903), not based on a generic 

Orders Pyrsonymphina, Oxymonadina, Retortomonadina, and Distomata Grasse 
Traite Zool. 1: fasc. 1: 788, 801, 824, 963 (1952). 
Non-pigmented flagellates with simple or acroneme flagella of definite number, 
from four to eight (two in Retortomonas), in the individual neuromotor system, and 
accordingly on the individual cell, except when the neuromotor systems are multi- 
plied; not of the definite characters of the following order. Free-living, chiefly in 
foul waters, or commensal or parasitic in animals. Polymastix is presumably the type 
of the group. It was listed with a query in Biitschli's original publication of family 

In the generality of Polymastigida, the cells are dorsiventral and have single nuclei 
and neuromotor systems. There are derived examples in which the cells are spirally 
twisted. There is a group in which the cells are double, having two nuclei and neuro- 
motor systems. In another group there are two or more neuromotor systems, usually 
with more than one nucleus; the cells consist of units in a whorled or spiral arrange- 
ment, so that as wholes they are of radial symmetry. 

The neuromotor system consists primarily of ( 1 ) the flagella; (2) one or more 
blepharoplasts from which the flagella spring; (3) one or more rhizoplasts linking 
together the parts of the system; and (4) a centrosome located just outside the nuclear 
membrane. Furthermore, (5) a parabasal body may be present. (6) An axostyle is a 
rod imbedded in the cytoplasm. In Hexamita the axostyles are the proximal ends of 
backwardly directed flagella; axostyles occurring in various other genera of the order 
appear also to be homologous with flagella. 

Nuclear and cell division have been observed in various genera, as in Hexamita by 
Swezy (1915); in Streblomastix by Kidder (1929); in Giardia by Kofoid and Chris- 
tianson (1915) and Kofoid and Swezy (1922); and in O.V);mona^ by Connell (1930). 
Cleveland (1947) observed in Saccinobaculus a multiplication of nuclei followed 
by their fusion in pairs, and by meiosis in the fusion nuclei: thus there is a .sexual 
cycle without fusion of cells. It is not probable that sexual reproduction does not 
occur in the generality of the group, but it has not been observed in any others. 


The Classification of Lower Organisms 

Fig. 31. — PoLYMASTAcroA : a, Polymastix Mclolonthae after Swezy (1916). b, 
Streblomastix Strix x 1,000 after Kidder ( 1929) . c, d, Giardia cnterica after Kofoid 
& Swezy (1922). Trichomonadina: e, Hcxamastix Tcrmopsidis after Kirby 
(1930). i' Tricercomitus Termopsidis 2ihtv YJirhy (1930). g, Macrotrichomonas 
pulchra after Kirby (1938). h. Trichomonas tenax x 4,000 after Hinshaw (1926). 
i, Pentatrichomonas obliqua after Kirby (1943). j, Snydcrella Tabogae x 500 after 
Kirby ( 1929) . x 2,000 except as noted. 

Phylum Protoplasta [165 

In making the clearly natural group of trichomonads a separate order, Kirby ( 1947 ) 
removed the majority of the species formerly assigned to this order, and left a mis- 
cellany of small isolated families. It seems not expedient to make them several small 
orders, as Grasse has done; rather they are to be held together until their respective 
relationships become evident. A hint of Hall has led in the present work to the trans- 
fer of family Trimastigida to order Ochromonadalea. 
1. With a single nucleus and neuromotor system. 
2. Cells not spirally twisted, at least not as 

wholes and not conspicuously Family 1. TEXRAMiTroA. 

2. Entire cells conspicuously spirally 

3. With four free flagella Family 2. Streblomastigida. 

3. With four or eight flagella whose 
proximal ends are grown fast to the 

cell membrane Family 3. Dinenymphida. 

1. With one or several nuclei and two or more 

neuromotor systems Family 4. Oxymonadida. 

1. With two nuclei and neuromotor systems Family 5. Trepomonadida. 

Family 1. Tetramitida [Tetramitidae] Kent Man. Inf. 1: 312 (1880). Families 
Tetramitina and Polymastigina Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 841, 
842 (1884). Family Tetramitaceae Senn in Engler and Prantl Nat. Pflanzenfam. I 
Teil, Abt. la: 143 ( 1900) . Family Polymastigidae Doflein Protozoen 83 ( 1901 ) . Fam- 
ily Chilomastigidae Wenyon (1926). Family Costiidae Kudo Handb. Prot. 153 
(1931). Family Retortomonadidae Wenrich 1932. Cells mostly dorsiventral and 
with four flagella; these uniform or differentiated; when differentiated, one or two 
may trail behind the cell. Axostyles present or absent, parabasal bodies not reported. 
Like the order, the family is a miscellany; good authority has made as many as four 
families of the few genera. Tetramitus, free-living, unfamiliar. Costia, occurring 
usually as sessile parasites on fishes. Polymastix, in insects. Monocercomonoides, in 
insects and vertebrates. Chilomastix, in insects and vertebrates, cells marked by a 
cytostomal groove into which one of the flagella, shorter than the others, is recurved. 
The species which occurs in man (usually, as it appears, as a harmless commensal) 
is in most works called C. Mesnili; the correct name is apparently Chilomastix Hom- 
inis (Davaine) n. combl. Current authority places next to Chilomonas the biflagellate 
Retortomonas, also in insects and vertebrates, and having cells of essentially the 
same structure. 

^Kofoid (1920) gave the history involved in this combination. Davaine, 1860, de- 
scribed the flagellates Cercomonas Hominis var. A and var. B. The two forms are 
not of the same species, and Moquin-Tandon, in the same year, re-named them 
respectively C. Davainei and C. obliqua. They are not of the same genus, being re- 
spectively a Chilomastix and a Pentatrichomonas, under which genera they have 
various names. Kofoid named them respectively Chilomastix davainei and Tricho- 
monas hominis. In so doing, he may be held to have exercised his right to choose a 
type in a group in which no type has been designated; but it is arguable on the con- 
trary that an author who designates a var. A designates the type in doing so. It is 
on the basis of this argument that the new combination here published is applied 
to the Cercomonas Hominis var. A of Davaine. 

166] The Classification of Lower Organisms 

Family 2. Streblomastigida [Streblomastigidae] Kofoid and Swezy in Univ. Cali- 
fornia Publ. Zool. 20: 15 (1919). The only known species is Strehlornastix Strix, a 
slender spirally twisted organism with four anterior flagella, free-swimming or at- 
tached in the gut of the termite Termopsis. The significance of the epithet Strix (a 
Greek noun meaning screech owl) as applied to this species is not clear. 

Family 3. Dinenymphida [Dinenymphidae] Grassi in Atti Accad. Lincei ser. 5. 
Rendiconti CI. Sci. 20, 1° Semestre: 730 (1911). Elongate flagellates, the four or 
eight anterior flagella adherent to the body and spirally twisted with it, free at their 
distal ends. Often beset with spirochaets, which have been mistaken for additional 
flagella; the family has been misplaced in order Hypermastigina. Dinenympha and 
Pyrsonympha in termites; Saccinohaculus in the wood roach Cryptocercus. 

Family 4. Oxymonadida [Oxymonadidae] Kirby in Quart Jour. Micr. Sci. n. s. 72: 
380 ( 1928) . Flagellates with radially symmetrical bodies including two or more neuro- 
motor systems, entozoic in termites of subfamily Kalotermitinae. Each pear-shaped 
cell of Oxymonas has one nucleus and two neuromotor systems (Kofoid and Swezy, 
1926). In Microrhopalodina {Proboscoidella) each cell contains a whorl of nuclei, 
each with its separate neuromotor system (Kofoid and Swezy, 1926; Kirby, 1928). 
These organisms are superficially closely similar to the Calonymphida, from which 
Kirby distinguished them. 

Family 5. Trepomonadida [Trepomonadidae] Kent Man. Inf. 1: 300 (1880). 
Family Hexamitidae Kent op. cit. 318. Distomata Klebs in Zeit. wiss. Zool. 55: 329 
(1893). Family Distomataccae Senn in Engler and Prantl Nat. Pflanzenfam. I Teil, 
Abt. la: 148 (1900). Flagellates each with two nuclei and two neuromotor systems. 
In most examples, each half-cell is dorsiventral, and the whole isobilateral, with two 
cytostomes. Most of the genera, Trepomonas, Gyromonas, Trigonomonas, are free- 
living in fresh or foul waters and have been little studied. Hexamita occurs both free- 
living and entozoic, in roaches and in all classes of vertebrates; the cells have eight 
flagella {Octomitus Prowazek and Urophagus Moroff are synonyms). In Giardia 
the half-cells are asymmetric, and the whole cells dorsiventral, with one cytostome. 
There are several species, serious pathogens in mammals. The valid name of the 
species in man, usually known as G. Lamblia, appears to be G. enterica (Grassi) 
Kofoid (1920). 

Order 3. Trichomonadina Grasse Traite Zool. 1, fasc. 1: 704 (1952). 

Order Trichomonadida Kirby in Jour. Parasitol. 33: 215, 224 (1947), preoc- 
cupied by family TRiCHOMONADroAE Wenyon (1926). 

Flagellates of the general nature of the Polymastigida having in each neuromotor 
system one trailing flagellum; axostyle present, rigid, apparently not homologous 
with the flagella; parabasal body present, disappearing during mitosis. Entozoic, the 
majority of the species, to the number of fully 150, occurring in termites. 

The base of the trailing flagellum may be underlain by a cresta, a more or less 
prominent body distinct both from parabasal body and from axostyle. The trailing 
flagellum may be grown fast to the cell membrane and converted into an undulating 
membrane; in this case it is underlain by a rod called the costa, apparently homolo- 
gous with the cresta (Kirby, 1931). 

Nuclear and cell division have been described in Trichomonas by Kuczynski 
(1914), Kofoid and Swezy (1915, 1919; the Trichomitiis described in the latter 
year is a Trichomonas) and Hinshaw (1926). The centrosome (or a combined cen- 
trosome and blcpharoplast, the centroblcpharoplast of Kofoid and Swezy, 1919) lies 

Phylum Protoplasta [167 

outside the nuclear membrane. This structure divides and the daughter structures 
move apart along the nuclear membrane. They remain connected, usually until mito- 
sis is complete, by a stainable strand, the paradesmose. Definite chromosomes, usually 
few in number, and an intranuclear spindle, are formed. Mitosis is completed by con- 
striction of the nuclear membrane. In what appears to be the typical course of cell 
division, the rhizoplast and blepharoplast divide when the centrosome does. Of other 
parts of the neuromotor system, some may remain connected to one blepharoplast 
and some to the other; some may disappear. The parts needed to complete a neuro- 
motor system are regenerated in each daughter cell. 
1. With a single nucleus and neuromotor system. 
2. Lacking a cresta, costa, or undulating 

membrane Family 1. MoNOCERCOMONADroA. 

2. With a trailing flagellum whose base is 

underlain by a cresta Family 2. DEVEScoviNroA. 

2. With a trailing flagellum grown fast to 
the cell membrane, forming an undula- 
ting membrane underlain by a costa Family 3. Trichomonadida. 

1. With several nuclei and neuromotor systems. . Family 4. CALONYMPHroA. 
Family 1. Monocercomonadida [Monocercomonadidae] Kirby in Jour. Parasitol. 
33: 225 (1947). Minute flagellates of the appearance of certain Tetramitida, but 
having a firm axostyle, the parabasal body disappearing and a paradesmose forming 
between the daughter centrosomes during mitosis; lacking a cresta, costa, or undulat- 
ing membrane; entozoic in termites and other insects, and in all classes of vertebrates. 
Monocercomonas, Hexamastix, Tricercomitus. 

Family 2. Devescovinida [Devescovinidae] Doflein Lehrb. Prot. ed. 3: 537 (1911). 
Subfamily Devescovininae Kirby in Univ. California Publ. Zool. 36: 215 (1931). 
Organisms with three anterior flagella and a larger trailing flagellum underlain by a 
cresta; confined to termites of the families Mastotermitidae, Hodotermitidae, and 
Kalotermitidae, being most abundant in the last. The cells, usually fairly large, ingest 
scraps of wood and are presumed to contribute to the lives of their hosts by digesting 
it. Devescovina, Gigantomonas, Macrotrichomonas, Foaina, Parajoenia, Metadeves- 
covina. Spirochaets which share the habitat of these organisms are commonly found 
adhering to their cell membranes, and were mistaken for additional flagella in the 
original descriptions of some of the genera. 

Family 3. Trichomonadida [Trichomonadidae] Wenyon Protozoology 1 : 646 
(1926). Flagellates with three or more flagella directed forward and one trailing, the 
proximal part of the latter grown fast to the cell membrane and forming an undula- 
ting membrane underlain by a costa. Entozoic in a wide variety of animals. Tricho- 
monas, normally with four anterior flagella, is the most numerous genus. It occurs 
in termites, including those of the advanced family Termitidae, in which scarcely 
any other flagellates occur; it does not ingest wood, and is not believed to be benefi- 
cial to its hosts. It occurs also in all classes of vertebrates. Man harbors Trichomonas 
tenax as a commensal in the mouth. T. vaginalis may be a serious pathogen. Penta- 
trichomonas obliqua (Moquin-Tandon) comb. nov.,l commensal (or pathogenic?) 
in the gut has at the anterior end a fifth flagellum separate from the other four 

icf. footnote, p. 165. 

168] The Classification of Lower Organisms 

Family 4. Calonymphida [Calonymphidae] Grass! in Atti Accad. Lincei ser. 5, 
Rendiconti CI. Sci. 20, 1° Semestre: 730 (1911). Flagellates with radially symmetri- 
cal bodies including more than two nuclei and neuromotor systems, the latter of 
trichomonad type; entozoic in termites of subfamily Kalotermitinae. These flagellates 
ingest scraps of wood and are believed to contribute to the nutrition of their hosts. 
In Coronympha each cell contains one whorl of nuclei each with its separate neuro- 
motor system (Kirby, 1929). In Stephanonympha, the nuclei and neuromotor systems 
are so numerous as to form a spiral band of several cycles in the anterior part of the 
cell. In Calonympha, besides numerous neuromotor systems associated with nuclei, 
there are others free of any nucleus; in Snyderella, the two types of structures are 
independently multiplied. 

Order 4. Hypermastigina Grassi in Atti Accad. Lincei ser. 5, Rendiconti CI. Sci. 
20, 1° Semestre: 727 (1911). 
Order Trichonyynphidea Poche in Arch. Prot. 30: 149 (1913). 
Order Hypermastigida Calkins Biol. Prot. 29"5 (1926). 

Order Lophomonadida Light in Univ. California Publ. Zool. 29: 486 (1927). 
Orders Joeniidca, Lophomonadina, Trichonymphina, and Spiratrichonym- 
phina, Grasse Traite Zool. 1, fasc. 1: 837, 851, 862, 916 (1952). 

Flagellates, mostly large and of radial symmetry, with single nuclei and indefi- 
nitely numerous flagella. Entozoic in roaches and in termites excluding those of 
family Termitidae. Lophomonas is to be regarded as the type. 

Cleveland (1925, 1926) found it possible, by starvation or by exposure to high 
pressures of oxygen or high temperatures, to rid insects of all of their intestinal 
flagellates or of some of the kinds. When completely freed of flagellates, wood roaches 
and termites of the lower families are able to remain alive only for a few weeks. The 
life of Termopsis is not prolonged by the presence of Streblomastix, and it is pro- 
longed only moderately by the presence of Trichomonas Termopsidis. But if infested 
with either Trichonympha Campanula or T. sphaerica, it can survive indefinitely on 
a diet of pure cellulose. Both species ingest the ground scraps of wood which reach 
the part of the intestine in which they occur; it is evident that they serve their hosts 
as agents of digestion. Cleveland's observations raise unanswered questions as to the 
occurrence of fixation of nitrogen; it is known only that termites are quite economical 
in their use of nitrogenous compounds available to them. 

The Hypermastigina have elaborate neuromotor systems. There is regularly a large 
centroblepharoplast. In what appears to be the relatively primitive type of cell divi- 
sion, as in Trichonympha (Kofoid and Swezy, 1919), the neuromotor system of the 
mother cell is divided between the daughter cells. In Spirotrichonympha (Cupp, 
1930), only the centroblepharoplast divides; the neuromotor system of the mother 
cell remains attached to one of the daughter centroblcpharoplasts, while the other 
generates the remaining parts of a complete system. In Lophomonas (Kudo, 1926), 
and Kofoidia (Light, 1927), the neuromotor system of a dividing cell is absorbed 
or discarded, with the exception of the centroblcpharoplasts, from which new systems 

In Trichonympha and Spirotrichonympha the details of nuclear division have 
much the appearance of meiosis. A double set of chromosomes appears, and the 
chromosomes form pairs which are divided in the spindle. It is supposed that this 
appearance is produced by a precocious splitting of the chromosomes. 

Phylum Protoplasta [ 169 

In species of Trichonympha, Leptospironympha, and Eucomonympha from the 
wood roach Cryptocercus, Cleveland (1947, 1948) observed the syngamy of undiffer- 
entiated or diflFerentiated gametes; the appearance of the process is as though the egg 
ingested the sperm. Syngamy is followed immediately by meiosis. This means that 
vegetative individuals are haploid. Barhulanympha achieves without syngamy an al- 
ternation of haploid and diploid stages. Diploid cells are produced when a centro- 
blepharoplast fails to divide, with the result that the nucleus remains intact, while 
chromosomes appear and divide. Reduction division, by the separation of undivided 
chromosomes, occurs when a centroblepharoplast divides at an exceptionally early 
stage. Cleveland concluded that the early division of the central body is the event 
which primarily distinguishes meiosis from mitosis. It is possible that he has recog- 
nized an essential feature of the evolution of the sexual cycle. His words suggest the 
idea that the sexual cycle may have originated within the present group. This is an 
impossibility; the sexual cycle is a normal character of nucleate organisms, and is 
fully established in nucleate organisms far more primitive than these. 

There are fewer than one hundred known species of Hypermastigina. They are 
treated as seven families. 

1. Body without segmented appearance. 

2. Flagella distributed generally over the 

surface of the body or its anterior part. . . . Family 1. TRiCHONYMPHroA. 

2. Flagella in spiral bands Family 2. HoLOMASTiGOTororoA. 

2. Flagella in tufts. 

3. Flagella in a single tuft Family 3. LoPHOMONAoroA. 

3. Flagella in two tufts Family 4. HoPLONYMPHroA. 

3. Flagella in four tufts Family 5. SxAUROjOENnDA. 

3. Flagella in many tufts Family 6. KoForonoA. 

1. Body with segmented appearance Family 7. Teratonymphida. 

Family 1. Trichonymphida [Trichonymphidae] Leidy ex Doflein Lehrb. Prot. ed. 
3: 537 (1911). The numerous flagella distributed generally over the surface of the 
body or its anterior part. Trichonympha {Leidy opsis), Eucomonympha, etc. 

Family 2. Holomastigotoidida [Holomastigotoididae] Janicki in Zeit. wiss. Zool. 
112: 644 (1915). Family S pirotrichonymphidae Grassi in Mem. Accad. Lincei CI. 
Sci. ser. 5, 12: 333 (1917). The numerous flagella arranged in spiral bands. Holo- 
m.astigotoides, S pirotrichonympha, etc. 

Family 3. Lophomonadida [Lophomonadidae] Kent Man. Inf. 1: 321 (1880). 
Family Joeniidae Janicki in Zeit wiss. Zool. 112: 644 (1915). The numerous flagella 
assembled in a single anterior tuft. Lophomonas, in cockroaches, all of the flagella 
directed forward. Joenia, Joenina, Joenopsis, etc., in termites, the outer flagella 
directed backward. 

Family 4. Hoplonymphida [Hoplonymphidae] Light in Univ. California Publ. 
Zool. 29: 138 (1926). The flagella assembled in two anterior tufts. Hoplonympha, 
Barhulanympha, etc. 

Family 5. Staurojoeninda [Staurojoenindae] Grassi in Mem. Accad. Lincei CI. Sci. 
ser. 5, 12: 333 (1917). The flagella assembled in four anterior tufts. Staurojoenina. 

Family 6. Kofoidiida [Kofoidiidae] Light in Univ. California Publ. Zool. 29: 485 
(1927). The flagella fused at their bases into several bundles. Kofoidia, a single 
known species in Kalotermes. 

Family 7. Teratonymphida [Teratonymphidae] Koidzumi in Parasitology 13: 303 
(1921). Family Cyclonymphidae Reichenow. Elongate and segmented, with a single 


The Classification of Lower Organisms 

Fig. 32. — Hypermastigina : a-d, Trichonympha Campanula after Kofoid & 
Swezy (1919); a, cell x 250; b, division of centroblcpharoplast and formation of 
paradesmose, and c and d, earlier and later stages of mitosis x 500. e, f, g, Sperm, 
egg, and fertilization of Trichonympha sp. from the roach Cryptocercus after Cleve- 
land (1948). h, Hoplonympha Natator x 250 after Light (1926). i, Staurojoenina 
assimilis x 250 after Kirby (1926). j, Tcratonympha mirabilis after Koidzumi 

Phylum Protoplasta [171 

nucleus in the anterior segment; flagella distributed generally on the surface, most 
abundant on an anterior beak. Teratonympha Koidzumi {Cyclonympha Dogiel), a 
single known species in Reticulitermes. 

Class 2. MYCETOZOA de Bary 

Order Dermatocarpi Persoon Syst. Meth. Fung, xiii (1801), in part. 

Suborder Myxogastres Fries Syst. Myc. 3: 3 (1829); suborder Trichospermi Fries 

op. cit. 1 : xlix (1832), in part. 
Suborder MyATomyce^^j Link 1833. 

Mycetozoen de Bary in Bot. Zeit. 16: 369 (1858); Zeit. wiss. Zool. 10: 88 (1859). 
Stamm Myxomycetes Ylatcktl Gen. Morph. 2: xxvi (1866). 
Class Mycetozoa de Bary ex Rostafinski Versuch Systems Mycetozoen 1 (1873). 
Division Mycetozoa and classes Myxogasteres and Phytomyxini Engler and Prantl 

Nat. Pflanzenfam. IITeil: 1 (1888). 
Division Myxothallophyta Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, 

Abt. 1: iii (1897). 
Stamm Myxophyta Wettstein Handb. syst. Bot. 1: 49 (1901). 
Division Phytosarcodina, Myxothallophyta, or Myxomycetes Engler Syllab. ed. 3: 

1 (1903). 
Division Myxomycophyta Tippo in Chron. Bot. 7: 205 (1942). 
Order Mycetozoida Hall Protozoology 227 (1953). 

Organisms whose walled resting cells produce in germination anteriorly unequally 
biflagellate cells; these giving rise to bodies called plasmodia, being multinucleate 
bodies of amoeboid character. 

1. Predatory, subaerial, producing macroscopic 

spore-bearing fruits. 

2. Spores produced within the fruits Order 1. Enteridiea. 

2. Spores produced on the surfaces of the 

fruits Order 2. Exosporea 

1. Parasitic, not producing definite fruits Order 3. Phytomy.xii>a. 

Order 1. Enteridiea [Enteridieae] Rostafinski Vers. 3 (1873). 

Cohort Endosporeae and orders Anemeae, Heterodermeae, Reticularieae, Ain- 
aurochaeteae, Calcareae, and Calonemeae Rostafinski op. cit. 

Order Endosporea Lankester in Enc. Brit. ed. 9, 19: 840 (1885). 

Orders Protodermieae and Columniferae Rostafinski ex Berlese in Saccard) 
Sylloge7: 328,417 (1888). 

Cohorts Amaurosporales and Lamprosporales, with numerous orders with names 
in -aceae. Lister Monog. Mycetozoa 21-23 (1894). 

Subclass Myxogastres and orders Physaraceae, Stemonitaceae, Cribrariaceae, 
Lycogalaceae, and Trichiaceae Macbride North American Slime Molds 20 

Subsuborder (!) Endosporinei Poche in Arch. Prot. 30: 200 (1913). 

Orders Physarales, Stemcnitales, Cribrariales, Lycogalales, and Trichiales Mac- 
bride op. cit. cd. 2 (1922). 

Order Liceales Ma. bride and Martin (1934). 

Suborder Eumycetozoina Hall Protozoology 230 (1953). 

172 ] The Classification of Lower Organisms 

Predatory Mycetozoa producing macroscopic fruits, these producing internal uni- 
nucleate spores. The type is Lycogala, the sole genus of the order as originally 

The fruits of many examples are of the appearance of minute puffballs, and Per- 
soon and Fries classified them as puffballs; Fries took note that they are primitus 
mucilaginosi and made them a suborder distinct from the proper pufTballs. De Bary 
studied the non-reproductive stages; concluded "dass die Myxomyceten nicht dem 
Pflanzenreiche angehoren, sondern dass sie Thiere, und zwar der Abtheilung der 
Rhizopoden angehorig, sind"; and renamed the group Mycetozoen. This name was 
apparently first published in Latin form, in the category of classes, by de Bary's stu- 
dent Rostafinski. Conventional botany continues to list Myxomycetes as a class of 
Fungi; conventional zoology makes the group an order of Rhizopoda or Sarcodina. 

The spores germinate readily in water or appropriate solutions (Jahn, 1905; Gil- 
bert, 1929; Smith, 1929). Their nuclei usually divide once or twice, during or just 
after germination; thus each spore produces from one to four naked cells. 

It is in germinating spores that mitosis is most easily observed. Mitosis takes place 
in a clear area, about which some observers have found a persistent nuclear membrane. 
The spindle is sharp-pointed. Only a few observers (as Skupienski, 1927) have dis- 
cerned definite centrosomes. When the one or two divisions associated with germina- 
tion are complete, the flagella grow forth from the areas of the poles of the mitotic 
spindle. All earlier observers described the spores as uniflagellate, but Ellison (1945) 
and Elliott (1949) found them biflagellate. The flagella may be apparently equal or 
moderately unequal; or one of them may be very brief. Each nucleus remains con- 
nected to the base of the flagella by a conical body of clear protoplasm, the Geissel- 
glocke of Jahn (Jahn, 1904; Howard, 1931). 

The flagellate cells are not spores, but gametes; they fuse with each other. Skupien- 
ski (1917) affirms that they are of two mating types. Fusion is at first by pairs, and 
Howard (1931) found that each zygote develops into a plasmodium by itself. All 
other observers (de Bary, 1858, 1859; Cienkowski, 1863; Skupienski, 1917, 1927; 
Schiinemann, 1930) have found the zygotes to fuse with each other and with further 
gametes. The flagella are lost. The nuclei fuse in pairs; those which fail to find 
partners are digested. 

The cell formed by the fusion of zygotes and gametes is a young plasmodium. The 
term was coined by Cienkowski ( 1863, p. 326) : "Das Protoplasmanetz der Myxomy- 
ceten werde ich mit den Namen Plasmodium bezeichnen." The plasmodium nour- 
ishes itself in predatory fashion, on fungus spores, bacteria, and other digestible ob- 
jects, and grows accordingly. Mitosis occurs simultaneously in all nuclei of the plas- 
modium, and takes 20 to 40 minutes; it has accordingly only rarely been observed 
(Lister, 1893; Howard, 1932). Plasmodia do not ordinarily divide, but grow to great 
sizes. They are not very familiar objects because during most of their life they keep to 
dark and moist places, chiefly among vegetable remains. Drouth does not kill them; 
they can become dry and hard while retaining the capacity to resume activity upon 
the return of moisture. When an active plasmodium reaches a certain stage, its re- 
actions change; it moves out into the light and to dry places. A plasmodium in this 
stage is conspicuous, being of the form of a network which may be many centimeters 
in diameter, in some species brilliantly colored. The whole is a single naked protoplast. 

Each Plasmodium proceeds to produce a fruit or fruits. The entire mass may heap 
itself up, or it may break up into portions, large or minute. In species whose plas- 
modia break up into small fragments, each of these may secrete a column of lifeless 

Phylum Protoplasta [173 

material, a millimeter or more in height, and ascend upon it. Each separate body of 
protoplasm secretes an external wall and begins to undergo cleavage within it. Har- 
per (1900) described the details of the process. All authorities agree that the nuclei 
undergo a flare of divisions at this time (Strasburger, 1884; Harper, 1900, 1914; 
Bisby, 1914). It is almost certain that there are two flares of division, constituting the 
meiotic process, but few authorities have positively affirmed this (Schiinemann, 
1930)1. Cleavage is carried to the point of producing uninucleate protoplasts. While 
this is taking place, many species secrete a network of hollow tubes or a system of 
hollow fibers, called the capillitium, by deposition of lifeless material outside the 
cell membranes. In species which produce a true capillitium, all of the uninucleate 
protoplasts secrete walls and become spores. Strasburger found the capillitium and 
the walls of the spores to consist of impure cellulose; others have found no cellulose. 
In many species which do not produce a true capillitium, an analogous structure 
called a pseudocapillitium, consisting of solid bodies of various forms, is modelled 
from a part of the nucleate protoplasm which is deprived of its reproductive function 
and killed. In many species, much calcium carbonate is deposited in the wall, or 
both in the wall and in the capillitium or pseudocapillitium. 

A small separate fruit is called a sporangium. A fruit of the form of a large mass, 
or of many sporangia not completely separate, is an aethalium. The spores are re- 
leased by collapse of the outer wall. 

These organisms are of no known economic importance. There are some forty 
genera, between four and five hundred species. As Lister remarked, the same species 
occur everywhere: collections from Colombia (Martin, 1938) and from Mount 
Shasta (Cooke, 1949) consist entirely of familiar species. 

Rostafinski (1873) arranged the genera in two cohorts, seven orders, and nineteen 
tribes, the last with names in -aceae. His subsequent monograph of the group ( 1875) 
was regrettably published in a barbarous language, and is for nomenclatural purposes 
a nullity. All later systems are based on Rostafinski's original system. The group being 
essentially uniform, it is properly treated as a single order. 

Definite families were first established by Lankester, mostly under names which 
Rostafinski had applied to tribes. Berlese (in Saccardo, 1888) provided a complete 
set of names in -aceae, valid under botanical rules; Poche provided a complete set in 
-idae, valid under zoological rules. Authorities have differed moderately as to the 
list of families; here, somewhat arbitrarily, fourteen are maintained. 
1. Capillitium none (order Cribrariales Mac- 

2. Producing separate sporangia, pseudo- 
capillitium none. 

3. Sporangia shattering irregularly or 
opening through a terminal oper- 
culum Family 1 . Liceacea. 

3. Sporangia opening through numer- 
ous pores, the walls becoming sieve- 
like Family 2. Cribrariacea. 

2. Fruits aethalioid, pseudocapillitium 

iWhile the present work was in proof, Wilson and Ross (1955) established the point 
that meiosis occurs immediately before the formation of spores. 

174 ] The Classification of Lower Organisms 

3. Aethalia consisting of more or less 
separate sporangia. 

4. Sporangia tubular, opening 

through terminal pores Family 3, Tubiferida. 

4. Sporangia indistinct, their walls 
becoming freely punctured and 
converted into a reticulate 

pseudocapillitium Family 4. Retigulariacea. 

3. Aethalia not consisting of distin- 
guishable sporangia Family 5. Lycogalacttoa. 

1. Capillitium present. 

2. Fruits without considerable deposits of 
calcium carbonate. 

3. Spores black or dark, capillitial 
hairs smooth (order Stemonitales 

4. Fruits aethalioid, capilHtium 
poorly defined, without a cen- 
tral axis Family 6. Amaurochaetacea. 

4. Fruits of separate sporangia 
with a definite capillitium in- 
cluding a central axis (colu- 

5. Capillitium spreading hor- 
izontally from the colu- 
mella Family 7. Stemonitea. 

5. Capillitium spreading 
chiefly from the summit of 

the columella Family 8. Enerthenemea. 

3. Spores pallid or yellow (order Tri- 
chiales Macbride). 

4. Capillitial hairs smooth, un- 
branched or sparsely branched. 
5. Capillitial threads hori- 
zontal, attached at both 

ends Family 9. Margaritida. 

5. Capillitial threads run- 
ning at random, not at- 
tached at the ends Family 10. Perichaenacea. 

4. Capillitium reticulate, sculp- 
tured, but not with spiral bands. . .Family 11. Arcyriagea. 
4. Capillitial threads unbranched 
or sparsely branched, sculp- 
tured with spiral bands Family 12. Trighiagea. 

2. Fruits containing considerable deposits 
of calcium carbonate (order Physarales 

3. Calcium carbonate both in walls 

and in capillitium Family 13. Physarea. 

Phylum Protoplasta [175 

3. Calcium carbonate in walls but not 

in capillitium Family 14. DrovMiACEA. 

Family 1. Liceacea [Liceaceae] (Rostafinski) Lankester in Enc. Brit. ed. 9, 19: 
841 (1885). Tribe Liceaceae Rostafinski Vers. 4 (1873). Order Liceaceae Lister 
Monog. Mycetozoa 149 (1894). Family Liceidae Doflein 1909. Family Orcadel- 
lidae Poche in Arch. Prot. 30: 200 (1913). Family Orcadellaceae Macbride N. Am. 
Slime Molds ed. 2: 203 (1922). Sporangia separate, sessile or stalked, without capil- 
litium or pseudocapillitium, the walls shattering irregularly or opening by means of 
a terminal operculum. Licea, Orcadella. 

Family 2. Cribrariacea [Cribrariaceae] (Rostafinski) Lankester 1. c. Tribe Crib- 
rariaccac Rostafinski op. cit. 5. Order Cribrariaceae Macbride N. Am. Slime Molds 
20 (1899). Order Heterodermaceae Lister op. cit. 136. Family Cribrariidae Poche 
1. c. The wall of the stalked fruit becoming sieve-like. Cribraria. Dictydium. 

Family 3. Tubiferida [Tubiferidae] Poche in Arch. Prot. 30: 200 (1913). Order 
Tubulinaceae Lister op. cit. 152 (1894). Family Tubulinidae Doflein 1909. Family 
Tubiferaceae Macbride in N. Am. Slime Molds ed. 2: 203 (1922). Aethalia consist- 
ing of tubular sporangia opening through terminal pores. Tubifer (its older name 
Tubulina preoccupied), Lindbladia, Alwisia. 

Family 4. Reticulariacea [Reticulariaceae] (Rostafinski) Lankester 1. c. Tribes 
Dictydiaethaliaceae and Reticulariaceae Rostafinski op. cit. 5, 6. Order Reticularia- 
ceae Lister op. cit. 156. Family Dictydiaethaliidae Poche I.e. Aethalia of indistinct 
sporangia whose walls become porous and are converted into a reticulate pseudo- 
capillitium. Reticularia, Dictydiaethallium, etc. 

Family 5. Lycogalactida [Lycogalactidae] Poche in Arch. Prot. 30: 201 (1913). 
Tribe Lycogalaceae de Bary. Order Lycogalaceae Macbride N. Am. Slime Molds 20 
(1899). Y di.m.i\y Lycogalaceae Macbride and Martin Myxomycetes (1934). Aethalia 
with a pseudocapillitium, not divided into sporangia. Lycogala, the brownish fruits a 
few millimeters in diameter clustered on wood, of much the appearance of small 

Family 6. Amaurochaetacea [Amaurochaetaceae] (Rostafinski) Berlese in Sac- 
cardo Sylloge 7: 401 (1888). Tribe Amaurochaetaceae Rostafinski op. cit. 8. Order 
Amaurochaetaceae Lister op. cit. 134. Family Amaurochaetidae Doflein 1909. 
Fruits aethalioid with dark spores and a poorly defined capillitium without a central 
axis. Amaurochaete. 

Family 7. Stemonitea Lankester in Enc. Brit. ed. 9, 19: 841 (1885). Tribes Stemo- 
nitaceae and Brefeldiaceae Rostafinski op. cit. 6, 8. Families Stemonitaceae and 
Brefeldiaceae Berlese in Saccardo op. cit. 390, 402. Order Stemonitaceae Macbride 
N. Am. Slime Molds 20 (1899). Family Stemonitidae Doflein 1909. Families Bre- 
feldiidae and Stemonitidae Poche op. cit. 202. Sporangia with dark spores and a 
capillitium of smooth threads spreading from a central axis, the columella. Stemo- 
nitis, comm.on, the clustered stalked fruits of the appearance of minuscule dark 
bottle-brushes. Brefeldia, Comatricha; Diachea, exceptional in containing much lime 
in the stalk and wall. 

Family 8. Enerthenemea Lankester 1. c. Tribes Echinosteliaceae and Enerthene- 
maceae Rostafinski op. cit. 7, 8. Families Echinosteliaceae and Enerthenemaceae 
Berlese in Saccardo op. cit. 389, 402. Family Lamprodermaceae Macbride N. Am. 
Slime Molds ed. 2: 189 (1922). Like Stemonitea, in which this family has usually 
been included, but the capillitium attached chiefly at the summit of the columella. 
Enerthenema, Clastoderma, Lamproderma, Echinostelium. 


The Classification of Lower Organisms 

Family 9. Margaritida [Margaritidae] Doflein 1909. Order Margaritaceae Lister 
op. cit. 202. Family Dianemaceae Macbride N. Am. Slime Molds ed. 2: 237 (1922). 
Sporangia with pale or yellow spores and a capillitium of smooth threads attached at 
both ends. Dianema, Margarita. 

Family 10. Perichaenacea [Perichaenaceae] (Rostafinski) Lankester 1. c. Tribe 
Perichaenaceae Rostafinski op. cit. 15. Sporangia with pale or yellow spores and a 
capillitium of unattached smooth threads. Perichaena, Ophiotheca. 

Family 11. Arcyriacea [Arcyriaceae] (Rostafinski) Lankester 1. c. Tribe Arcyri- 
aceae Rostafinski op. cit. 15. Order Arcyriaceae Lister op. cit. 182. Family Arcyriidae 
Doflein 1909. Sporangia with pale or yellow spores and a reticulate capillitium, 
usually sculptured, but not with spiral bands. Arcyria, Lachnobolus. 

Family 12. Trichiacea [Trichiaceae] (Rostafinski) Berlese in Saccardo Sylloge 7: 
437 (1888). Tribe Trichiaceae Rostafinski op. cit. 14. Family Trichinaceae Lankes- 

Fig. 33. — Mycetozoa. a-f, Spore, germination, gametes, syngamy, and zygote of 
Physarum polycephalum after Howard (1931) x 1,000. g-1, Stages of mitosis in the 
Plasmodium of Physarum polycephalum after Howard ( 1932) x 2,000. m-o. Stages 
of mitosis in the plasmodium of Trichia after Lister (1893) x 1,000. p, Cleavage 
in the developing fruit of Physarum polycephalum after Howard (1931) x 1,000. 
q, Capillitium and spores of Lepidoderma Chailletii x 1,000. r-W, fruits of Myce- 
toza X 5; r, Sternonitis splcndens; s, Lycogala cpidcndrum; i, Lcocarpus fragilis; 
U, Lepidoderma Chaillettii; v, Physarum notabile; w, Hemitrichia intorta. 

Phylum Protoplasta [177 

ter 1. c; the genus Trichina does not belong to this family! Order Trichiaceae Mac- 
bride N. Am. Slime Molds 20 (1899). Family Trichiidae Doflein 1909. Sporangia 
with pale or yellow spores, the capillitium of free threads, unbranched or sparsely 
branched, marked with spiral bands. Trichia, Hemitrichia, Oligonema, Calonema. 

Family 13. Physarea Lankester 1. c. Tribes Cienkowskiaceae, Physaraceae, and 
Spumariaceae Rostafinski op. cit. 9, 13. Families Cienkowskiaceae , Physaraceae, and 
Spumariaceae Berlese in Saccardo op. cit. 328, 329, 387. Order Physaraceae Macbride 
N. Am. Slime Molds 20 (1899). Family Physaridae Doflein 1909. Fruits sporangial 
or aethalioid, with capillitium, both wall and capillitium containing considerable 
deposits of calcium carbonate. Physarum, with some seventy-five species, is the most 
numerous genus of Mycetozoa; the little gray sporangia may be spherical or irregular, 
sessile or stalked. Fuligo septica produces dirty yellow aethalia reaching several cen- 
timeters in diameter on vegetable trash; observed on spent tan bark, it has the com- 
mon name of flowers of tan. Badhamia, Craterium, Leocarpus, Chondrioderma, 
Spumaria, etc. 

Family 14. Didymiacea [Didymiaceae] (Rostafinski) Lankester 1. c. Tribe Didy- 
miaceae Rostafinski op cit. 12. Order Didymiaceae Lister op. cit. 93. Family Didy- 
midae Doflein 1909. Family Didymiidae Poche op. cit. 202. Family Collodermata- 
ceae Macbride and Martin Myxomycetes 145 (1934). Sporangia with deposits of 
calcium carbonate in the wall and a simple capillitium free of mineral deposits. 
Didymium, Leangium, Lepidoderrna, Colloderma. 

Order 2. Exosporea (Rostafinski) Lankester in Enc. Brit. ed. 9, 19: 841 (1885). 

Cohors Exosporeae Rostafinski Vers. 2 (1873). 

OrAtr Ectosporeae\tr ?>y\\2ih. 2 (1892). 

Order Ceratiomyxaceae (Schroter) Lister Monog. Mycetozoa 25 (1894). 

Subsuborder (!) Exosporinei Poche in Arch. Prot. 30: 200 (1913). 
Organisms of much the character of the Enteridiea, but the spores forming a single 
layer on the surface of the fruits. There is a single family with only one well-marked 

Family Ceratiomyxacea [Ceratiomyxaceae] Schroter (in Engler and Prantl, 1889). 
Ceratiomyxa Schroter [Ceratium Albertini and Schweinitz, 1805, non Schrank, 1793); 
C. fruticulosa (O. F. Miiller) Macbride. The fruits are white pillars, sometimes 
branched, 1-2 mm. tall, of secreted material. Each spore of the single superficial 
layer generates a microscopic stalk and ascends upon it before becoming walled. 
Meiosis then takes place, making the spores 4-nucleate; the chromosome number is 
cut from 16 to 8 (Gilbert, 1935). In germination, the contents of the spore are re- 
leased as a single amoeboid protoplast, whose nuclei divide once; the cell then divides 
into eight, and these generate flagella (Rostafinski, 1873; Jahn, 1905; Gilbert, 1935). 

Order 3. Phytomyxida Calkins Biol. Prot. 330 (1926). 

Class Phytomyxini Engler and Prantl Nat. Pflanzenfam. II Teil: 1 (1889); class 

Phytomyxinae op. cit. I Teil, Abt. 1: iii (1897). 
Order Phytomyxinae Campbell Univ. Textb. Bot. 71 (1902). 
Class Plasmodiophorales Engler Syllab. ed. 3: 1 (1903). 
Order Plasmodiophorales Sparrow in Mycologia 34: 115 (1942). 
Suborder Plasmodia phorina Hall Protozoology 228 (1953). 
Intracellular parasites chiefly of higher plants, attacking also algae, Oomycetes, 
and beetles, being naked multicellular plasmodia producing walled resting cells, 


The Classification of Lower Organisms 

the walls containing no cellulose; these releasing naked infective colls with paired 
unequal simple flagella. 

This inconsiderable group was made known by the discovery of Plasmodiophora 
Brassicae, the agent of the clubroot disease of cabbage, by Woronin (1878). The 
proper place of the group in classification has been a puzzle; some students treat it 
as a class of myxomycetes, others as an order of chytrids. The known characters — 
paired unequal simple flagella; cells naked in the vegetative condition; and non-pro- 
duction of cellulose — assure us that this group has nothing to do with proper chytrids, 
nor with Oomycetes of chytrid body type. The traditional association with myxomy- 
cetes is tenable. Alternatively, the group would not be out of place next to order 
Rhizoflagellata (anyone who chooses to put it there should take note that the class 
name Phytomyxini is older than Zoomastigoda). 

The Plasmodium causes often much hypertrophy of the host tissue. In some forms 
the mature plasmodium becomes walled; the protoplast undergoes cleavage into uni- 
nucleate portions; these become swimming cells and are released through a discharge 
tube. These forms are of much the appearance of Lagenidialea. In the majority of 
the group the naked plasmodium undergoes cleavage; the resulting protoplasts be- 
come walled; the resulting spores or cysts, released by decay of the host, discharge 
their contents as one or two swimming cells. Ledingham ( 1939) and Sparrow ( 1947) 
report both types of development as occurring in Polyniyxa. Karling (1944) found 
the walls to contain no cellulose. Ellison (1945) found the flagella to be simple. 

iKyuiU/Tnui wuium/^iiiMiivnirm- 

Fig. 34. — Ceratiomyxa jruticulosa. a. Fruits x 5. b-q, reproductive processes 
after Gilbert (1935); b, young spores on the surfaces of the fruit; c, d, the same 
raised on stalks; e^ f, heterotypic division; g, homeotypic division; h, the mature spore 
ou its stalk; i-n, germination and subsequent processes: the amoeboid protoplast 
passes through a "thread stage" before rounding up and dividing into four and then 
into eight; o, production of flagcllum; p, "zoospore" (gamete); q, gametes fusing to 
initiate the plasmodium. All x 1,000 except Fig. a. 

Phylum Protoplasta [179 

In the growing plasmodium, a nucleus which is not dividing contains an endosome 
("nucleolus"). During mitosis, which occurs within the intact nuclear membrane, 
the endosome becomes elongate, and a ring of chromatin, within which separate 
chromosomes have not been distinguished, forms about its middle. The resulting "cru- 
ciform" figure resembles some which have been seen in trypanosomes. The nuclear 
divisions which occur immediately before cleavage are of a different character: no en- 
dosome is seen, but there is a spindle with centrosomes at the poles, and definite 
chromosomes are present. The occurrence of these two types of nuclear division has 
been noted by every careful observer, Schwartz (1914), Home (1930), Cook (1933), 
Ledingham (1939), and Karling (1944). Home was probably correct in supposing 
the divisions which precede cleavage to be meiotic. Conjugation of the flagellate cells 
of Spongospora has been observed. 

There are monographic accounts of the Phytomyxida by Cook ( 1933) and Karling 
(1942). The group may be treated as a single family with a dozen genera and about 
twenty-five species. 

Family Plasmodiophorea [Plasmodiophoreae] Berlese in Saccardo Sylloge 7 : 464 
(1888). Family Plasmodiophoreen Zopf Pilzthiere 129 (1885). Family Plasmodio- 
phoraceae Engler Syllab. 1 (1892). Family Woroninaceae Minden 1911. Families 
Phytomyxidae and Woroninidae Poche in Arch Prot. 30: 198 (1913). Plasmodio- 
phora, Polymyxa, Spongospora, and Sorosphaera attack land plants; Tetramyxa, 
Ligniera, and Sorodiscus, chiefly aquatic seed plants; Woronina and Octomyxa, 
Oomycetes; Phagomyxa, brown algae; Sporomyxa (Leger, 1908) and Mycetosporid- 
ium, beetles. 

Class 3. RHSZOPODA Siebold 

Order Foraminiferes d' Orbigny in Ann. Sci. Nat. 7: 128, 245 (1826). 

Order Foraminifera Zborewski 1834. 

Rhizopodes Dujardin in Compt. Rend. 1: 338 (1835). 

Class Foraminifera d'Orbigny in de la Sagra Hist. Cuba vol. 8 (1839). 

Order Polythalamia Ehrenberg in Abh. Akad. Wiss. Berlin (1838) : table 1 ( 1839) . 

Class Rhizopoda and orders Monosomatia and Polysomata Siebold in Siebold 
and Stannius Lehrb. vergl. Anat. 1 : 3, 11 (1848). 

Reticulosa Carpenter 1862. 

Stamm Rhizopoda and Class Acyttaria Haeckel Gen. Morph. 2: xxvii (1866). 

Thalamophora R. Hertwig Hist.Radiolar. 82 (1876). 

Class Reticidaria Lankester in Enc. Brit. ed. 9, 19: 845 (1885). 

Order Reticulosa Poche in Arch. Prot. 30: 203 (1913). 

Order Granuloreticulosa de Sacdeleer in Mem. Mus. Roy. Hist. Nat. Belgique 60: 
7 (1934). 

Order Foraminiferida Hall Protozoology 250 (1953). 

Amoeboid organisms, the pseudopodia of the character of rhizopodia, i.e., fine, 
freely branching and anastomosing; producing shells, these usually calcareous; com- 
monly reaching macroscopic dimensions; mostly marine. 

The first examples of rhizopodes mentioned by Dujardin were milioles, vorticiales, 
and le gromia: the genus Miliola is to be construed as the type. These organisms, the 
proper rhizopods, are in general usage called Foraminifera, but that name was orig- 
inally applied in the categoiy of orders. 


The Classification of Lower Organisms 

Fig. 35. — Life cycle of "Tretomphalus," i.e., Discorbis or Cymbalopora, from 
Myers (1943); 1-3, microspheric individuals, in 3 releasing young megalospheric 
individuals; 4-8 megalospheric individuals; 9-12, gametes and syngamy. 

Phylum Protoplast a [181 

The individual rhizopod originates as a minute amoeboid cell which secretes a 
shell from which the pseudopodia project. In the fresh-water forms, each protoplast, 
after moderate growth, divides into two, one of which retains the original shell while 
the other secretes a new one. In some of the marine forms, the original protoplast, 
having a cylindrical or irregular shell, enlarges this as it grows. In the great majority 
of the group, the original shell, called the proloculus, is of definite size and form and 
has a constricted orifice. When the protoplast reaches a certain stage, it expands, pro- 
trudes from the orifice, and secretes an extension of the shell in the form of a second 
chamber. In some few examples, the second chamber is the final one, being capable 
of indefinite extension. But again in the great majority, the second chamber, although 
diff'erent from the proloculus, resembles it in being definite in form and in having a 
constricted orifice. After further development, the protoplast again protrudes through 
the orifice and secretes a third chamber, generally of the same form as the second, 
though often larger. Repetition of this process produces macroscopically visible 
bodies. Even though becoming a centimeter or more in diameter, the individuals 
continue to be single cells. 

As a result of different patterns of growth, the developed shells are of highly varied 
forms, linear, globular, or coiled in one plane; trochoid or rotaloid, that is, helical, 
of the form of a low cone; of the form of high cones; or screw-like, with the chambers 
in fixed longitudinal rows. The grov/th pattern may change during the life of the 
individual. There are apparently degenerate forms, simple or irregular. It is highly 
probable that some of the forms have evolved repeatedly. 

The shells may be of gelatinous material or of chitin, without or with imbedded 
grains of sand. Exceptionally, they are siliceous. They are sometimes of crystallized 
calcium carbonate with imbedded grains of sand. In the bulk of the group they consist 
of crystallized calcium carbonate without foreign matter, and are of either of two 
t>'pes of texture: vitreous, that is, hyaline, and punctured by numerous pores a few 
microns in diameter; or porcellanous, white by reflected light and amber by trans- 
mitted light, and with no perforations except the proper orifices. In fossil shells, other 
textures than these may occur; it is supposed that these are products of modification 
during preservation. Some of the textures, like some of the forms, are believed to 
have evolved repeatedly. 

Most rhizopods occur in two forms which are most readily distinguished by the 
size of the proloculi. This was first pointed out by Munier-Calmas, 1880; who, 
jointly with Schlumberger, 1885, designated the smaller and larger proloculi re- 
spectively microsperes and megaspheres. Lister (1895), by study in culture of Elphi- 
dium crispiirn [Polystomella crispa Lamarck), showed that the two forms are alter- 
nate generations. He observed that the microspheric cells become multinucleate 
during growth, while the megalospheric cells remain uninucleate until just before 
reproduction. The reproduction of the megalospheric cells is by release of numerous 
minute biflagellate cells. 

Schaudinn (1902) confirmed much of what Lister had observed. He was mistaken 
in describing nuclear division (except just before the production of the swimming 
cells) as non-mitotic; and correct in identifying the swimming cells as gametes. 
Winter (1907) observed a similar life cycle in Peneroplis, but described the gametes 
as having solitary flagella. 

Myers" (1934, 1935, 1936), dealing with Patellina and Spirillina, described the 
details of mitosis. This takes place within an intact nuclear membrane, and is com- 
pleted by its constriction. The spindle is blunt-ended; there is no evidence of centre- 

182 ] The Classification of Lower Organisms 

somes. The chromosomes are numerous, long, and slender; the mitotic figures re- 
semble those of Pyrrhophyta. Reduction of the chromosome number is said to be 
effected by a single nuclear division, the last one before the formation of gametes, 
which cuts the chromosome number of Patellina from 24 to 12, and that of Spirillina 
from 12 to 6. Before they reach this stage, the megalospheric individuals have 
gathered themselves in clusters of two or more within cyst walls consisting of secreted 
gelatinous matter and scraps from the neighborhood. Gametes from one individual 
are unable to unite with each other. The gametes are amoeboid, positively without 
flagella. In Discorbis and Cymbalopora, however, Myers (1943) observed the produc- 
tion of biflagellate gametes. 

Le Calvez (1950) has cleared up various questions raised by earlier studies. Some 
forms, as Discorbis orbicularis, appear to lack a sexual cycle. Patellina and Spirillina 
produce amoeboid gametes 40-50[.i in diameter. Most rhizopods produce biflagellate 
gametes 1.5-4[i long. Le Calvez found the flagella definitely unequal. In Discorbis 
mediterranensis he showed that the megalospheric individuals are of two mating 
types. Earlier zoologists, apparently misled by familiarity with the normal life cycle 
cf animals, had identified meiosis as occurring at the time of gametogenesis; it is the 
fact, on the contrary, that it occurs in the last two nuclear divisions in the micro- 
spheric individuals. The megalospheric and microspheric stages of rhizopods are 
respectively haploid and diploid, like the gametophytes and sporophytes of plants. 

With the possible exception of some of the one-chambered fresh water forms, the 
rhizopods are clearly a natural group. The fresh water forms appear to intergrade 
with organisms which Pascher identified as chrysomonads. 

The shells of dead rhizopods may under appropriate conditions be preserved 
through geologic ages. Natural chalk consists of shells of Textularia mixed with coc- 
coliths. Certain forms of limestone consist chiefly of shells of Miliola. Certain fossil 
rhizopods have long been known as indicators of division of geologic time. Since about 
1917, it has been found that the whole group offers one of the beautiful illustrations 
of evolution as related to geologic time: the shells of rhizopods found under magnifi- 
cation in a particular stratum serve promptly and precisely to identify it. The services 
of experts on "Foraminifera" have acquired a high economic value in the petroleum 
industry: these experts have found themselves promoted from the status of pure 
biologists to that of economic geologists. 

Among some eleven hundred genera which have been published, Galloway (1933) 
maintains 542. Of the number of species one can only say that it is a matter of 
thousands, but probably not many tens of thousands. Economic micropaleontologists 
find themselves dealing with great numbers of forms which are slightly, yet signifi- 
cantly, distinct. They find it expedient not to name these, but to identify them by 
comparison with available collections. 

Some of the marine and fossil forms are similar, on a small scale, to the animal 
Nautilus, and Linnaeus placed some of them in that genus. Montfort and Lamarck 
treated them as several genera of mollusks. In first distinguishing these organisms as 
the order Foraminiferes of class Cephalopodes, d'Orbigny intended to contrast them 
with Nautilus, in whose shells a series of chambers arc connected, not by holes (fora- 
mina) but by cylindrical tubes. Dujardin ( 1835) found that his Rhizopodes are with- 
out definite organs. Their shells enclose a clear semiliquid substance; their apparent 
tentacles are merely temporary structures, formed of this substance, thrust forward 
in the direction of the movement of the shell and withdrawn as it advances. Dujardin 
named this substance sarcode; it is, of course, the same which has since been called 

Phylum Protoplasta [ 183 

protoplasm. The effect of his discoveries was to show that the rhizopods or Foramini- 
fera are not mollusks, but one-celled organisms. 

Very much taxonomic study has been given to this interesting group. The standard 
system, in the modern period of practical concern with the group, has been that of 
Cushman (1928). 

Galloway (1933), attempting to recognize phylogeny and concluding that certain 
types of form and texture of shells have evolved repeatedly, has radically revised 
Cushman's system and set up a system of thirty-five families. The following survey 
of the group is based on Galloway's system. The names applied to the families are 
those which he has cited as the oldest, and the groups treated as orders are the blocks 
of families which to him appeared natural. 

1. Shell one-chambered, or of a proloculus fol- 
lowed by one other chamber, not of a series 

of similar chambers Order 1. Monosomatia. 

1. Shell a series of similar chambers. 

2. Shell porcellanous, imperforate Order 2. Miliolidea. 

2. Not as above. 

3. Not specialized as in the following 

orders Order 3. Foraminifera. 

3. Shell hyaline, perforate, typically 
trochoid, i.e., having the succes- 
sively larger chambers helically ar- 
ranged so that all may be seen from 
one side and only the last whorl 

from the other Order 4. GLOBiGERiNroEA. 

3. Chambers of the fundamentally 
planispiral shell with specialized 
walls containing channels or pro- 
ducing chamberlets Order 5. Nummulitinidea. 

Order 1. Monosomatia (Ehrenberg) Siebold in Siebold and Stannius Lehrb. vergl. 

Anat. 1: 11 (1848). 
Monosomatia Ehrenberg in Abh. Akad. Wiss. Berhn (1838) : table 1 (1839). 
Order Astrorhizidea Lankester in Enc. Brit. ed. 9. 19: 846 (1885). 
Order Imperforida Delage and Herouard Traite Zool. 1: 107 (1896). 
Order Archi-Monothalamia Calkins Biol. Prot. 354 ( 1926) . 
Rhizopoda consisting of a single chamber, or of a proloculus followed by one other 
chamber; exceptionally, after passing through a stage of this character, producing 
a series of similar chambers. 

Family 1. Allogromiida [Allogromiidae] Cash and Wailes. Minute, with one- 
chambered chitinous or gelatinous shells, usually subglobular; large in fresh water. 
Allogromia Rhumbler; Mikrogromia Hertwig, the pseudopods of sister cells retaining 
contact so that small colonies are formed; etc. 

Family 2. Astrorhizida [Astorhizidae] Brady (1881). Family Astrorhizina Lankes- 
ter (1885). Family Astrorhizidaceae Lister. Families Rhizamminidae , Saccammini- 
dae, and Hyperamminidae Cushman. Shell of agglutinated foreign material, usually 
elongate, often branched, but not coiled. In Astrorhiza there is a central chamber 
from which grow elongate arms. In Rhizammina, the shell is tubular, open at both 
ends; in Bathysiphon it is a tube closed at one end; in Hyperammina a proloculus is 
formed before the extended tube. 


The Classification of Lower Organisms 

- ^ / m\^■^V^//'":^^^^^.^V^\\■•\\ 

Fig. 36 — Shells of Rhizopoda. a, Ophthalimidium. h, c, Triloculina. d, Verte- 

bralina. e, Peneroplis. f, Archaias x 25. g, Nodosaria. h, Dcntilina. i, Flabel- 

lina. j, Lagena. k, 1, Nonion. m, n, Rotalia. o, Globigerina. x 50 except as 

Phylum Protoplasta [ 185 

Family 3. Spirillinidea Reuss 1861. Family Spirillinina Lankester (1885). Family 
Silicinidae Cushman. In Spirillina, the perforate hyaline one-chambered shell is 
planispirally coiled; the family is distinguished by a shell of this form in the young 
stages if not throughout life. Silicina is a Jurassic fossil whose shell is silicified. In 
Patellina the spirally coiled first chamber is followed by others arranged in a heHx. 

Family 4. Ammodiscida [Ammodiscidae] Rhumbler 1895. Like the preceding 
family, but the shell consisting of agglutinated foreign matter. Ammodiscus etc. 

Order 2. Miliolidea Lankester in Enc. Brit. ed. 9, 19: 846 (1885). 
Order Flexostylida Calkins Biol. Prot. 355 (1926). 

Rhizopoda with imperforate porcellanous shells, a numerous and important group. 

Family 1. Miliolida [Miliolidae] d'Orbigny (1839). Families N uhecularina, Milio- 
lina, and Hauerinina Lankester (1885). Fisherinidae Cushman. The genus Cornu- 
spira, known from the carboniferous, differs from Spirillina only in the texture. Evi- 
dently evolved from this are genera of planispirally coiled tubes divided into chambers, 
and from these others in which the series of chambers becomes straight or irregular, 
as in Vertebralina and Tubinella. There is an important block of genera in which each 
cycle of chambers is of two members, the second opening at the opposite end of the 
body from the first. In O phthalmidium and Pyrgo alternate chambers lie regularly 
on opposite sides of a body whose form is that of an elliptic flake. In other genera 
of this group successive chambers are not opposite each other, but separated by less 
than 180°, so that more than two appear on the outside. In Triloculina three cham- 
bers are externally visible. In Miliola Lamarck [Miliolina Lamarck, the latter name 
applied to fossil representatives of the same genus) the chambers are 144° apart, so 
that five appear on the outside. In many members of the family the apertures are 
partially blocked by teeth, single, double, or multiple, or extended as bars clear across. 

Family 2. Soritina Ehrenberg (1839). Family Helicosorina Ehrenberg op. cit. 
Family Peneroplidea Reuss 1861. Family Peneroplidina Lankester (1885). Family 
Peneroplidae Cushman. Family Soritidae Galloway (1933). Specialized derivatives 
of the lower Miliolida: planispiral shells in which the chambers become successively 
larger, as in Peneroplis, and, by a further development, divided into large numbers 
of secondary chambers, as in Archaias, Sorites, and Orbitolites. Spirolina, the shell 
coiled in the oldest part, straight in the remainder. 

Family 3. Alveolinea Ehrenberg (1839). Family Alveolinida Schultze 1854. 
Families Alveolinina and Keramosphaerina Lankester, Alveolinellidae and Keramo- 
sphaeridae Cushman. Another group of specialized derivatives, the planispirally coil- 
ing chambers broadened and divided into many chamberlets with separate apertures, 
the entire body more or less globular. Borelis; Fasciolites Parkinson 1811 [Alveo- 
lina d'Orbigny 1826); Alveolinella; Keramosphaera Brody, a rare antarctic form. 
The organisms of these last two families resemble, as a parallel development, those 
of order Nummulitinidea, from which they are distinguished by the texture. 

Order 3. Foraminifera Zborewski 1834. 

Order Polythalamia and subordinate group Polysomatia Ehrenberg in Abh. 

Akad. Wiss. Berlin (1838): table 1 (1839). 
Order Polysomatia Siebold in Siebold and Stannius Lehrb. vergl. Anat. 1 : 11 

Orders Lituolidea, Textularidea, and Lagenidea Lankester in Enc. Brit. ed. 9, 

19: 847 (1885). 

186] The Classification of Lower Organisms 

Order Perforida Delage and Herouard Traite Zool. 1 : 107 (1896). 
Orders Nodosalida and Textulinida Calkins Biol. Prot. 355, 356 (1926). 

Comparatively unspecialized Rhizopoda, the shells of various textures, not porcel- 
lanous; not usually of trochoid form, and if so, usually not vitreous. 

Early students, Montfort, Lamarck, and d'Orbigny, were much concerned with 
organisms which they called Geophonus, Vorticialis, or Polystornella. These names 
represent organisms of much the appearance of Nautilus; all are synonyms of Elphi- 
dium Montfort, which is to be considered the type or standard genus of Foraminifera. 

Family 1. Endothyrina Lankester (1885). Family Endothyridae Rhumbler 1895. 
Fossils, pre-Cambrian to Carboniferous, the calcareous shells granular or fibrous, not 
porcellanous or vitreous. Cayeuxina Galloway ( 1933) includes minute globular shells 
solitary or irregularly clustered, described by Cayeux, 1894, from the pre-Cambrian 
of Brittany; Matthewina Galloway includes Cambrian fossils of similar character. 
Endothyra and Cribrospira are Carboniferous forms, planispirally coiled; Tetrataxis 
produced trochoid shells. 

Family 2. Nodosinellida [Nodosinellidae] Rhumbler 1895. Shells like those of 
the Endothyrina or containing imbedded grains of sand, one-chambered or forming 
straight or curved, not coiled, rows. Mostly Carboniferous, rare as late as the Eocene. 
Archaelagena, Nodosinclla, Nodosaroum, Pedangia, etc. 

Family 3. Reophacida [Reophacidae] Cushman 1827. A small group of forms ap- 
parently degenerate from the foregoing, the chambers in straight, curved, or irregular 
series, walls chitinous or sandy; sometimes parasitic in other rhizopods. Reophax, etc., 
surviving to the present in cold deep water. 

Family 4. Trochamminida [Trochamminidae] Schwager 1877. Family Trocham- 
minina Lankester (1885). Family Plocapsilinidae Cushman. Cells planispiral or 
trochoid, becoming evolute or irregular; walls with imbedded grains of sand. Penn- 
sylvanian to recent, abundant only in the Cretaceous. Trochamniina, Plocapsilina, etc. 

Family 5. Lituolidea Reuss 1861. Family Lituolidae Brady (1881). Families 
Lituolina and Loftusiina Lankester. Family Lituolidaceae Lister. Families Loftu- 
siidae and Neusinidae Cushman. Shells spiral or becoming evolute or irregular, with 
walls of agglutinated siliceous or calcareous matter, the chambers subdivided as in 
order Nummulitinidea. Cyclammina, Lituola, Loftusia, Neusina, etc.; Mississippian 
to recent, most abundant in the Cretaceous and at present. 

Family 6. Orbitolinida [Orbitolinidae] Martin 1890. Specialized derivatives of 
the preceding family, walls agglutinated as in that group, the numerous chambers 
forming a conical or nearly circular body. Dictyoconus, Orbitolina, etc. Mesozoic and 

Family 7. Ataxophragmidea Schwager 1877. Families Valvulinidae and Vcr- 
neulinidae Cushman 1927. Family Ataxophragmidae Galloway (1933). Having 
walls of agglutinated material and allied to the preceding families; chambers of the 
shell tending to form an elongate, screw-like spiral. Valvulina, Ataxophragmium, 
Verneulina, etc.; since early Mesozoic, abundant in the present. 

Family 8. Textularina Ehrcnberg (1839). Family Textularidac d'Orbigny (1839). 
Family Textulariaccac Lister. Walls more or less agglutinated, the chambers usually 
in an elongate spiral with two members to a cycle, so that they form two series, the 
body as a whole tending to be wedge-shaped. Textularia, Cuneolina, Vulvulina, etc.; 
Ordovician to the present. 

Family 9. Nodosarina Ehrcnberg (1839). Family Nodosarida Schultzc 1854. 
Family Lagenidae Brady (1881). Family Lagenina Lankester (1885). Family Nodo- 

Phylum Protoplasta [ 187 

saridae Rhumbler. Family Lagenaceae Lister. Walls calcareous, hyaline, perforate; 
chambers planispiral in the earliest forms, becoming curved or straight in the major- 
ity; orifice ordinarily of radiating slits, becoming reduced to a single slit. A numerous 
group, Triassic to the present. Lenticulina Lamarck {Lenticulites Lamarck and 
Crist ellaria Lamarck are synonyms) is Naiitilus-Vikt. Hemicristellaria and Vaginulina 
resemble the sheath of a dagger; Flabellina and Frondicularia resemble fans; Glandu- 
lina is shaped like a jug. Nodosaria is like a row of enlarging beads. Lagena is a one- 
chambered form connected to Nodosaria by transitions, and evidently reduced, not 

Family 10. Polymorphinida [Polymorphinidae] d'Orbigny. Families Polymorphin- 
ina and Ramulinina Lankester ( 1885). Specialized irregular forms related to the pre- 
ceding, as indicated by orifices of the same character. Polymorphina, etc., present in 
the Mesozoic, abundant in the Cenozoic to the present. 

Family 11. Nonionidea [Noninideae] Reuss 1860. Family Polystomellina Lankester 
(1885). Family Hantkeninidae Cushman. Shells mostly nautiloid, that is, plani- 
spiral with successively larger chambers, a few of the highest trochoid; walls hyaline, 
perforate; aperture generally a transverse slit. N onion Montfort {Nonionina d'Or- 
bigny) and Elphidium Montfort [Geophonus Montfort, Vorticialis Lamarck, Poly- 
stomella Lamarck, the apparent type of Foraminifera) are simply nautiloid; Hant- 
kenina is ornamented with spines. Jurassic to the present. 

Order 4. Globigerinidea Lankester in Enc. Brit. ed. 9, 19: 847 (1885). 

Orders Rotalidea and Chilostomellida Lankester 1. c, both names having prev- 
ious use in the category of families. 
Order Rotalida Calkins Biol. Prot. 356 (1926). 

The main body of Rhizopoda with perforate hyaline shells, many-chambered, the 
chambers primitively of the trochoid arrangement. 

Family 1. Rotalina Ehrenberg (1839). Family Rotalidea Reuss 1861. Family 
Rotalidae Brady (1861). Family Rotalina Lankester. Family Rotaliaceae Lister. 
Families Globorotaliidae, Anomalinidae , and Planorbidinidae Cushman. A numerous 
family, including unspecialized forms, Globorotalia, Rotalia, etc., as well as degen- 
erate and irregular forms, Piano pulvinulina, etc., and moderately specialized ones 
with conical or disk-shaped bodies of numerous chambers, Cymbalopora, Planorbu- 
lina, etc. Triassic, rare; Jurassic to the present, common. 

Family 2. Acervulinida Schultze 1854. Family Rupertiidae Cushman. A small 
group of degenerate derivatives of the foregoing, the bodies attached, irregular, some- 
times reduced to one chamber. Rupertia, Acervulina, etc.. Cretaceous to the present. 

Family 3. Tinoporidea Schwager 1877. Family Calcarinidae Cushman. Another 
small group derived from Rotalina, the disk-shaped cells with a whorl of prominent 
spines. Calcarina, Tinoporus, etc. Cenozoic, to the present. 

Family 4. Asterigerinida [Asterigerinidae] d'Orbigny (1839). Two genera, Asteri- 
gerina and Amphistegina, diverging from Rotalina in having each chamber divided 
into two by an oblique wall. Doubtfully in the Cretaceous; Eocene to the present. 

Family 5. Chapmaniida [Chapmaniidae] Galloway (1933). The numerous cham- 
bers arranged in a low cone whose inside is filled with deposited solid material. Chap- 
mania, Halkyardia, Dictyoconoides. Eocene and Oligocene. 

Family 6. Chilostomellida [Chilostomellidae] Brady (1881). Family Chilostomell- 
aceae Lister. A few genera of reduced derivatives of Rotalina with few chambers. 
Allomorphina, Chilostomella, Sphaeroidina, etc. Jurassic to the present. 

188 ] The Classification of Lower Organisms 

Family 7. Orbulinida Schultze 1854. Family Globigerinida Carpenter 1862. 
A few genera with the chambers mostly few, subglobular, clustered rather than ar- 
ranged in a definite pattern. Orbulina. Globigerina, abundant, pelagic in all oceans, 
the shells abundant in the ooze on the bottom. Pennsylvanian, doubtful; Jurassic, rare; 
Cretaceous to the present, common. 

Family 8. Pegidiida [Pegidiidae] Heron-Allen and Earland 1928. A few genera 
much like the Orbulinida but with thinner walls. Pegidia, etc. Oligocene to the 

Family 9. Heterohelicida [Heterohelicidae] Cushman 1927. A numerous group, 
the shells screw-like, biseriate, uniseriate, sheath-like or fan-like, the walls often with 
exterior ornamentation; paralleling the Nodosarina, but without the radiate orifices. 
Heterohelix, Sagrina, Eouvigerina, Pavonina, Plectojrondicidaria, Bolivina, Mucron- 
ina. Common, Jurassic to the present. 

Family 10. Buliminida Jones 1876. Family Uvellina Ehrenberg (1839), not 
based on a generic name. Family Buliminina Lankester (1885). Shells mostly high 
spirals, screw-like, often with spines or other external ornamentation, the orifices 
various, commonly comma-shaped. Turrilina, Bulimina, Virgulina, etc. Triassic to 
the present. 

Family 11. Cassidulinida [Cassidulinidae] d'Orbigny (1839). A small group with 
high-spiralled shells and comma-shaped orifices, evidently derived from the forego- 
ing family. Cassidulina, etc. Eocene to the present. 

Family 12 Uvigerinida [Uvigerinidae] Galloway and Wissler, 1927. Further vari- 
ants from Heterohelicida, the high-spiralled shells with chambers in three rows at 
first, varying to biseriate and uniseriate. Uvigerina, Siphonogenerina, etc. Jurassic 
to the present, common since the Miocene. 

Family 13. Pleurostomellida [Pleurostomellidae] Reuss 1860. An additional 
rather small family of the same general character as the few preceding. Pleurosto- 
mella, Nodosarella, Daucina, Ellipsoidina, etc. Cretaceous to the present, commonest 
in upper Cretaceous and Eocene. 

Order 5. Nummulitinidea Lankester in Enc. Brit. ed. 9, 19: 848 (1885). 

Rhizopoda with large specialized shells, the walls hyaline, perforate, generally 
thickened and traversed by channels and thrown into internal ridges which subdivide 
the chambers. 

Family 1. Fusulinida [Fusulinidae] Moller 1878. Carboniferous fossils, the 
chambers short and broad, numerous, in a planispiral coil, forming bodies which are 
usually fusiform or globular. Orobias, Fusidina, Triticina, Verbeekina, etc. 

Family 2. Nummulitida [Nummulitidae] Reuss 1861. Family Camerinidae 
Meek and Hayden 1865. Family Nummulinidae Brady (1881). Family Nummuli- 
tina Lankester ( 1885). Family Nummulitaceae Lister. Mostly disk-shaped, planispiral, 
the walls not highly specialized. Camerina Bruguiere 1792 {Nxunrnulites Lamarck 
1801), Operculina, Heterostegina, etc. Jurassic to the present, most abundant in the 

Family 3. Orbitoidida [Orbitoididae] Schubert 1920. Similar to the foregoing, 
the numerous chambers divided into numerous chamberlcts. A considerable group of 
Mcsozoic and Ccnozoic fossils. Orbitoides, Cyclosiphon, etc. 

Family 4. Cycloclypeina Lankester (1885). Family Cycloclypeidae Galloway 
(1933). Similar to the preceding. A number of Mcsozoic and Ccnozoic genera, most 
numerous in the Eocene. Asterocydina. The only living species is Cycloclypeiis Car- 
penteri Brady. 

Phylum Protoplasta [ 189 

Class 4. HEUOZOA Haeckel 

Family Polycystina Ehrenberg in Abh. Akad. Wiss. Berlin 1838: 128 (1839). 

Rhizopoda radiaria seu Radiolaria J. Miiller in Abh. Akad. Wiss. Berlin (1858) : 
16 (1859). 

Echinocystida Claparede. 

Order Radiolaria Haeckel Radiolarien 243 (1862). 

Stamm Moneres for the most part, and classes Heliozoa and Radiolaria, Haeckel 
Gen. Morph. 2: xxii, xxviii, xxix (1866). 

Subclasses Heliozoa and Radiolaria Biitschli in Brown Kl. u. Ord. Thierreichs 1, 1 
Teil: Inhalt (1882). 

Class Proteomyxa Lankester in Enc. Brit. ed. 9, 19: 839 (1885). 

Subclasses Proteomyxiae, Heliozoariae, and Radiolariae Delage and Herouard 
Traite Zool. 1: 66, 156, 169 (1896). 

Class Actinopoda Calkins Biol. Prot. 318 (1926). 

Class Actinopodea and orders Helioflagellida, Heliozoida, Radiolarida, and Proteo- 
myxida HaU Protozoology 202, 203, 212, 220 (1953). 

Subphylum Actinopoda Grasse and Deflandre, and classes Acantharia, Radiolaria, 
and Heliozoa Tregouboff in Grasse Traite Zool. 1, fasc. 2: 267, 270, 321, 437 

Organisms having pseudopodia of the character of filopodia, stiffly radiating, or of 
axopodia, stiffly radiating and having inner fibers; often with siliceous skeletons. 

Here, not without authority, one combines in one class the three groups which have 
been treated as the classes Proteomyxa, Heliozoa, and Radiolaria; and adds further 
two families of shelled amoebas. 

Cienkowski (1865) listed as "Monaden" the new species or genera Monas amyli, 
Colpodella (apparently a chytrid), Pseudospora, and Vampyrella. They are minute 
fresh-water amoeboid organisms, in part having flagellate stages. Haeckel (1866) 
placed most of them (the Monas under the new generic name Protomonas), together 
with his own discoveries Protamoeba and Protogenes, and also the bacteria, in his 
Stamm Moneres, i.e., his group of Protista without nuclei. Later (1868) he omitted 
the bacteria. Zopf (1885) found several of Haeckel's Moneres to possess nuclei, and 
Lankester renamed the group Proteomyxa. Publication of subsequent original obser- 
vations of these organisms has been scant and scattered; they remain poorly known. 

The Heliozoa as conventionally construed are also mostly inhabitants of fresh 
water. Ehrenberg observed some of them and took them for Infusoria with immobile 
cilia. There are only a few dozen species of Heliozoa sensu strict o (Schaudinn, 1896) : 
the whole group is no more than a reasonable order. 

The Radiolaria (this name also used at this point in its conventional sense) are 
marine. Examples were first observed as floating gelatinous bodies. These were taken 
for fragments and remained unnamed until 1834, when Mayen named Physematium 
and Sphaerozoum. Fossil skeletons of many examples were described by Ehrenberg 
(1839). Huxley (1851) named Thalassicolla and gave an accurate account of its 
structure. It was by work on organisms of this group that Haeckel first distinguished 
himself (1862). 

Haeckel dealt further with this group in four important papers (1879, 1882, 1887, 
1887-1888). In the last of these, the Radiolaria are a class of four legions, eight sub- 
legions, twenty orders, 85 families, 739 genera, and more than four thousand species. 
The categories, Haeckel explained, are purely relative: Radiolaria would as well be 

190] The Classification of Lower Organisms 

a phylum, the legions classes, and so forth. This idea served him as license for con- 
founding the application of many names, by shifting them among the categories, or 
by substituting new names for old. All subsequent authors have followed Haeckel's 
system of Radiolaria, applying names as best they might. 

The class Heliozoa in the extended sense here proposed may be organized as five 
orders distinguished as follows: 

1. Cells without a central capsule, i.e., without 
a firm membrane surrounding the inner part 

of the protoplast Order 1. Radioflagellata. 

1. Cells with a central capsule. 

2. Central capsule of spherical symmetry 
or with three planes of symmetry at 
right angles, punctured by many pores. 
3. Pores of the central capsule evenly 
distributed; skeleton absent or pres- 
ent, without spicules which cross the 

central capsule or meet in its center Order 2. Radiolaria. 

3. Pores of the central capsule clust- 
ered; skeleton including spicules 
which cross the central capsule or 

meet in its middle Order 3. Acantharia. 

2. Central capsule of radial symmetry, with 

one opening Order 4. Monopylaria. 

2. Central capsule of isobilateral symmetry, 
with one main opening and two minor 
ones Order 5. Ph.'SlEosphaeria. 

Order 1. Radioflagellata Kent Man. Inf. 1 : 225 ( 1880) . 

Subdivision or subclass Heliozoa (Haeckel), and orders Aphrothoraca (Hert- 

wig), Chlamydophora (Archer), Desmothoraca (Hertwig and Lesser), and 

Chalarothoraca (Hertwig and Lesser) Biischli in Bronn Kl. u. Ord. Thier- 

reichs 1: 261, 320, et seq. (1881, 1882). 

Suborder Protoplasta Filosa Leidy in Rept. U.S. Geol. Survey Territories 12: 

Class Proteomyxa Lankester ( 1885) . 

Subclass Proteomyxiae and orders Acystosporidia, Azoosporidia, and Zoospori- 
dia; subclass Heliozoariae and orders Aphrothoracida, Chlamydophorida, 
Chalarothoracida, and Desmothoracida Delage and Herouard Traite Zool. 
1: 66-72, 156-168 (1896). 
Order Heliozoa Doflcin Protozoen 13 ( 1901 ) . 

Orders Vampyrellidea and Chlamydomyxidea Poche in Arch. Prot. 30: 182, 
193 (1913). 
The proper Heliozoa together with the Proteomyxa: organisms of the character 
of the class, lacking central capsules, that is, firm membranes about the inner part of 
the protoplasts. Mostly fresh water organisms of spherical .symmetry, commonly with- 
out skeletons. The type, being the only genus assigned to the order by Kent, is 

1. Pscudopodia unspecialized; amoeboid organ- 
isms with or without flagellate stages. 

Phylum Protoplasta [191 

2. Without shells Family 1. Pseudosporea. 

2. With shells; without known flagellate 

3. Shells chitinous, without siliceous 

scales Family 2. Lagyntoa. 

3. Shells bearing circular siliceous 

scales Family 3. Euglyphida. 

1. Pseudopodia slender, with apical knobs Family 4. Vampyrellacea. 

1. Pseudopodia of the character of typical axo- 
podia, without apical knobs; the cells or their 
main bodies usually regularly spherical. 
2. Bearing flagella as well as axopodia in 

the vegetative condition Family 5. AcTiNOMONADroA. 

2. Without flagella in the vegetative 

3. Cells without a lifeless outer coat Family 6. AcTiNOPHRYroA. 

3. Cells having a gelatinous outer coat 

without siliceous spicules Family 7. Heterophryida. 

3. Cells having a gelatinous outer coat 

with siliceous spicules Family 8. Acanthocystida. 

3. Cells with a hard shell punctured 

by pores Family 9. Clathrulinida. 

Family 1. Pseudosporea [Pseudosporeae] Berlese in Saccardo Sylloge 7: 460 
fl888). Monadineae Zoosporcae Cienkowski in Arch. mikr. Anat. 1: 213 (1865). 
Family Pseudosporeen Zopf Pilzthiere 115 (1885). Orders Azoosporidea for the 
most part and Zoosporidca Delage and Herouard (1896). Azoosporidae for the most 
part and Zoosporidae Doflein Protozoen 40, 41 ( 1901 ) . Family Pseudosporidae Poche 
in Arch. Prot. 30: 197 (1913). Amoeboid organisms without shells or skeletons, the 
pseudopodia tapering from a broad base to a filamentous termination. Flagellate 
stages (with one flagellum or two unequal flagella) occur in Protovionas, Pseudo- 
spora. and Diplophysalis. In other genera, as Arachnula and Chlamydomyxa, no 
flagellate stages are known. 

Family 2. Lagynida Schultze 1854. Order Gromida Claparede and Lachmann 
1859. Family Gromida Carpenter 1862. Family Gromiidae Brady (1881). Families 
Monostomina and A.mphistomina Lankester (1885). Amoeboid organisms having 
chitinous shells without siliceous scales with a broad orifice through which project 
pseudopodia of the character of filopodia. Grom.ia, Lagynis, etc. 

Family 3. Euglyphida [Euglyphidae] Wallich 1874. Amoeboid organisms with a 
chitinous shell beset with circular siliceous scales, the filopodia projecting through a 
broad orifice. Euglypha, Cyphoderia, Campuscus, Trinema, etc. 

Family 4. Vampyrellacea [Vampyrellaceae] Zopf Pilzthiere 99 (1885). Monadin- 
eae Tetraplasteae Cienkowski op. cit. 218. Family Vampyrelleae Berlese in Saccardo 
Sylloge 7: 454 (1888). Family Vampyrellidae Poche in Arch. Prot. 30: 182 (1913). 
Cells subglobular, slowly creeping, with slender pseudopodia, numerous, densely 
packed and stiffly radiating on mature individuals, bearing terminal knobs. Vampy- 
rclla, the cells colored faintly pink by some metabolic by-product, is not unfamiliar 
as a predator on freshwater algae cultured under unfavorable conditions. 

Family 5. Actinomonadlda [Actinomonadidae] Kent Man. Inf. 1: 226 (1880). 
Family Ciliophryidae Poche in Arch Prot. 30: 187 (1913) Family Helioflagellidae 


The Classification of Lower Organisms 




h \ 

Fig. 37. — Radioflagellata : a-f, Diplophysalis stagnalis after Karling (1930); 
a, b, young cells with one or two flagella; c, active amoeboid form; d, walled cell; 
e, iame releasing flagellate cells; f, resting cell, g. Young cell of Vampyrella x 1,000. 
h, Actinosphaerium Eichhornii x 1,000. 

Phylum Protoplasta [ 193 

Doflein. Organisms bearing at the same time flagella and typical axopodia. Dimor- 
pha, free-swimming, with two unequal flagella. Actinomonas, Pteridomonas, Cilio- 
phrys, with one flagellum, either free-swimming or attached by a protoplasmic stalk. 
Family 6. Actinophryida [Actinophryidae] Glaus 1874. Askeleta Hertwig and 
Lesser in Arch. mikr. Anat. 10 Suppl. 164. (1874). Aphrothoraca seu Actinophryidae 
Hertwig Org. Radiolar. 142 (1879). Order Aphrothoraca Butschli (1881). Suborder 
Aphrothoraca Minchin (1912). Family Camptonematidae Poche in Arch. Prot. 30: 
187 (1913). Cells typically spherical, with typical axopodia, having no flagella nor 
shells nor skeletons. Actinophrys Sol Ehrenberg and Actinosphaerium Eichhornii 
(Ehrenberg) Stein are common in fresh water among algae, living as predators 
largely on diatoms; Actinophrys is uninucleate, the cells to 50^ in diameter; Actono- 
spaerium is multinucleate, the cells to lOOOjl in diameter. Camptonema is marine. 

Mitosis in Actinophrys as described by Schaudinn (1896) occurs within an intact 
nuclear membrane which undergoes constriction; the dividing nucleus lies within a 
spindle-like body of cytoplasm. Schaudinn and Belar (1923) observed conjugation. 
Pairs of gametes, which are usually sister cells but may sometimes be random pairs, 
lie within a cyst wall of secreted material. The nucleus of each gamete undergoes 
meiosis; at the end of each meiotic division, one of the daughter nuclei is digested; 
thus each gamete comes to possess a single haploid nucleus. Syngamy and karyogamy 
follow in due course and the zygote becomes walled. An old account of the cytology 
of Actinosphaerium by Hertwig is defaced by descriptions of the origin of nuclei from 
fragments of nuclei (chromidia), and of nuclear fusions at two separate stages of 

Family 7. Heterophryida [Heterophryidae] Poche in Arch. Pr«it. 30: 189 (1913). 
Heliozoa Chtamydophora Archer in Quart. Jour. Micr. Sci. n.s. 16: 348 (1876). 
Order Chlamydophora Butschli (1882). Suborder Chlamydophora Minchin (1912). 
Family Lithocollidae Poche I.e. The cells or their main bodies spherical with axopodia 
projecting through a gelatinous envelope. Heterophrys and Astrodisculus are simply 
globular cells. Elaeorhanis and Lithocolla are similar but with grains of sand or dia- 
tom shells imbedded in the envelope. Actinolophus is stalked. Sphaerastrum becomes 
colonial by incomplete division of the cells. 

Family 8. Acanthocystida [Acanthocystidae] Glaus 1874. Chalarothoraca Hert- 
wig and Lesser in Arch. mikr. Anat. 10 Suppl. 193 ( 1874). Chalarothoraca seu Acan- 
thocystidae Hertwig Org. Radiolar. 142. (1879). Order Chalarothoraca Butschli in 
Bronn Kl. u. Ord. Thierreichs 1: 325 (1882). Suborder Chalarothoraca Minchin 
(1912). Resembling the preceding family, but the gelatinous envelope containing 
hard bodies, supposedly usually siliceous, of definite form. In Raphidophrys, these 
bodies are curved needles; in Pinacocystis, small plates; in Acanthocystis and Pinacio- 
phora, disks bearing a central spine which is in some species forked. The cell of Wag- 
nerella (a marine form, on rocks in bays) consists of a globular head with spines and 
axopodia, borne on a protoplasmic stalk attached by a foot; the nucleus lies in the foot. 
In these forms the axial filaments of the pseudopodia radiate from a central gran- 
ule located outside the nucleus (in Wagnerella, in the head). Schaudinn (1896) re- 
ported nuclear division in Acanthocystis as being either amitotic or mitotic: the 
report of amitosis is of course not to be taken seriously. In the mitotic process, the 
central granule acts as a centrosome; the chromosomes are numerous and minute; 
the nuclear membrane disappears during the middle stages. Nuclear division may be 
followed by division of the cell into two, or may be repeated and followed by produc- 
tion of buds. The buds may lose their pseudopodia and develop paired flagella. It is 

194 ] The Classification of Lower Organisms 

suspected that the flagellate cells may be gametes. The central granule is said to 
originate by extrusion from the nucleus of a bud. 

Zuelzer (1909) found in Wagner ella two types of individuals, slender and stout, 
supposedly respectively haploid and diploid. In either type the nuclei may become 
numerous (and it is said that they sometimes develop from chromidia). The nuclei 
may migrate to the head and be released in buds, or they may become distributed 
throughout the protoplast, which then breaks up into biflagellate cells. It is supposed 
that these may be gametes, but a fusion of the heads of individuals of the slender 
type was observed. 

Family 9. Clathrulinida [Clathrulinidae] Glaus 1874. Desmothoraca Hertwig 
and Lesser op. cit. 225. Desmothoraca seu Clathrulinidae Hertwig Org. Radiolar. 
142 (1879). Order Desmothoraca Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 328 
(1882). Family Choanocystidae Poche in Arch. Prot. 30: 192 (1913). Protoplasts ly- 
ing within globular shells, apparently of chitin, usually stalked, punctured by numer- 
ous pores through which the axopodia project. In reproduction, the protoplast may 
divide into two, one of which escapes from the shell and secretes a new one; or it may 
divide into many which become unequally biflagellate. Clathrulina, Hedriocystis, 

Order 2. Radiolaria (J. Miiller) Haeckel Radiolarien 243 (1862). 

Rhizopoda radiaria sen Radiolaria J. Miiller in Abh. Akad. Wiss. Berlin 1858: 

16 (1859). 
Orders Thalassicollen, Sphacrozoen, and Peripyleen Hertwig Org. Radiolar. 133 

(1879). * 
Orders Pcripylaria, Collodaria, Symbelaria, and Syncollaria Haeckel in Jen- 

aische Zeitschr. 15: 447, 469, 471, 472 (1882). 
Legion S pumellaria or Peripylea, with orders Collodaria and Sphaerellaria and 

seven suborders, Haeckel in Rept. Voy. Challenger Zool. 18: 5, 9 ( 1887) . 
Legion Spiimellaria, sublegions Collodaria and Sphaerellaria, and six orders, 

Haeckel Radiolarien 2: 87 (1887). 
Order Peripylida Delage and Herouard Traite Zool. 1 : 176 ( 1896). 
Suborder Peripylaria Minchin Protozoa 225 (1912). 
Order Sphaeridca Poche in Arch. Prot. 30: 206 (1913). 
Order Peripylea Calkins Biol. Prot. 343 (1926). 
Suborder Peripylea Kudo Handb. Protozool. 259 (1931). 
Suborder Pen'py/ma Hall Protozoology 218 (1953). 
This order and the three which follow, being the Radiolaria as conventionally con- 
strued, are unicellular marine organisms with axopodia, having within the protoplast 
a layer of organic material, variously punctured and of various types of symmetry, 
which separates the inner protoplasm from the outer. The central capsule consists of 
this layer (the central capsule membrane) and its contents, including the one or more 
nuclei of the cell. Imbedded in the protoplasm there is usually a skeleton, usually sili- 
ceous, various in structure and sometimes highly complicated. The outer cytoplasm 
is commonly inhabited by symbiotic cryptomonads in the resting condition (yellow 
cells, zooxanthellae), and sometimes contains masses of dark material, apparently 
debris extruded from the central capsule. The type of Radiolaria is evidently Thalas- 
sicolla; this genus was the first one described from living material, and was listed 
first by J. Miiller in the original publication of the name. 

The order which includes Thalassicolla, and to which the name Radiolaria is here 
restricted, is distinguished by uniformly distributed small punctures in the central 

Phylum Protoplasta [ 195 

capsule membrane and by the absence of skeletal spicules extending across the central 
capsule or meeting in its middle. Except during reproduction, each central capsule 
contains a single nucleus, but the cells of many examples are coenocytic, containing 
several or many central capsules. 

Brandt (1885, 1905) observed reproduction particularly as it occurs by the produc- 
tion of swimming cells by some of the coenocytic forms. The nucleus divides to pro- 
duce very many, and the intracapsular cytoplasm divides to produce uninucleate 
flagellate cells. In Collosphaera, all the nuclei are included in these cells. The cells 
are of two sizes, produced by different individuals, and are supposed to be gametes. In 
Sphaerozoum and its allies, some of the nuclei degenerate instead of being included 
in the swimming cells, of which two sizes are produced by single individuals. It ap- 
pears that the swimming cells have characteristically two unequal flagella, though 
many are found to have only one, and some produce a third appendage by which 
they can attach themselves. 

Haeckel listed thirty-two families in his legion Spumellaria. Other authors recog- 
nize about a dozen, including the following. 

Family ThalassicoUida Haeckel (1882). Thalassicollen J. Miiller (1859). Family 
Collida Haeckel (1862); there is no corresponding generic name. Order Collida 
Haeckel (1887). Globular forms with a single central capsule, skeleton none or of 
numerous small spicules. Thalassicolla, Physematium, Lampoxanthium, etc. 

Family Sphaerozoida Haeckel (1882). Family Collozoida Haeckel op. cit. Family 
Sphaeroidina Haeckel (1862); there is no corresponding generic name. Coenocytic, 
each cell with several nuclei in separate central capsules; skeleton none or of numer- 
ous small spicules. Sphaerozoum, the cells globular, to 1 mm. in diameter; Raphido- 
zoum, the cells elongate. 

Family CoIIosphaerida Haeckel (1862). Coenocytic, the spherical cell to 1 mm. in 
diameter, with several central capsules, each with an individual lattice-like skeleton. 

Family Haliommatina Ehrenberg 1847. Families Ethmosphaerida, Ommatida, 
and Cladococcida Haeckel ( 1862) . Family Sphaerida Haeckel ( 1882) . Order Sphaer- 
oidea, with six families, Haeckel '(1887). Globular, with small numbers of radiating 
main spicules, the main spicules bearing tangential branches which form a globular 
network of definite pattern, or, often, two or more concentric networks. Haliomma, 
Actinomma, Hexacontium, Cladococcus, and many other genera. 

Further families are of the character of the Haliommatina, but with the spherical 
symmetry modified by abbreviation or elongation of one or more axes: 

Family Spongurida Haeckel (1862). Order Prunoidea, with seven families, 
Haeckel (1887). Having one axis elongate. Spongurus, Pipetta, etc. 

Family Lithocyclidina Ehrenberg 1847. Family Discida Haeckel (1862). Order 
Discoidea, with six families, Haeckel ( 1887) . Having one axis shorter than the others. 
Lithocyclia, Staurocyclia, Heliodiscus, etc. 

Family Larcarida Haeckel (1887). Order Larcoidea, with this and seven other 
families, Haeckel (1887). The skeleton with three unequal axes, or spiral. Cenolar- 
cus, etc. 

Order 3. Acantharia Haeckel in Jenaische Zeitschr. 15: 465 (1882). 
Order Actipyleen Hertwig Org. Radiolar. 133 (1879). 

Legion Acantharia or Actipylea, orders Acanthometra and Acanthophracta, and 
seven suborders, Haeckel in Rept. Voy. Challenger Zool. vol. 18 (1887). 


The Classification of Lower Organisms 











Fig. 38. — Radiolaria: a, motile cells of Collosphaera Huxleyi after Brandt ( 1885). 
b. Skeleton of Haliomma capillaris after Haeckel ( 1862) . c, Skeleton of Actinomma 
Asteracanthion after Haeckel (1862). d. Skeleton of //f/zorfz,?cu5 P/zaco^wcu^ after 
Haeckel (1887). Acantharia: e, Skeleton of Dorataspis costata after Haeckel 
(1387). Monopylaria: f, Central capsule of Tridictyopus elegans after R. Hertwig 
f 1879) . g, Skeleton of Lithocircus productus after R. Hertwig, op. cit. h. Skeleton 
of Eucyrtidium carinatum after Haeckel (1862). Phaeosphaerl^: i. Typical cen- 
tral capsule after R. Hertwig, op. cit. 

Phylum Protoplasta [ 197 

Legion Actipyea or Acantharia, sublegions Acanthometra and Acanthophracta, 
and orders Actinellida, Acanthonida, Sphaerophracta, and Prunophracta 
Haeckel Radiolarien II Teil (1887). 
Order Actipylida Delage and Herouard Traite Zool. 1 : 204 ( 1896) . 
Suborder Acantharia Minchin Protozoa 256 (1912). 
Order Acanthometrida Poche in Arch. Prot. 30: 212 (1913). 
Order Actipylea Calkins Biol. Prot. 345 (1926). 
Suborder Actipylea Kudo Handb. Protozool. 216 (1931). 
Suborder Actipylina Hall Protozoology 216 (1953). 

In this group the central capsule membrane has many punctures arranged in 
clusters. The skeleton includes radiating spicules; in some examples these extend 
through the cell from side to side, passing through the central capsule; in the majority, 
their proximal ends meet in the center of the central capsule. In the latter forms, 
the number of radiating spicules is twenty, and they are arranged according to a pat- 
tern discovered by Johannes Miiller and called Miiller's law; they form five parallel 
whorls of four. Usually they bear tangential branches of definite, often highly elab- 
orate, patterns: these may form a globular frame, or two or more concentric frames. 

Haeckel, Hertwig, and Brandt found the spicules not to consist of silica. They are 
soluble in acids and alkalis, and were reported to be destroyed by heat. They were 
supposed, accordingly, to consist of some organic substance; Haeckel named it acan- 
thin. Schewiakoff found them resistant to heat, and Biitschli (1906) analyzed them 
and found them to consist of strontium sulfate. This surprising fact has recently been 
confirmed by Odum (1951). 

The cytoplasm at each point where a spicule passes through the surface is attached 
to the spicule by a whorl of minute fibers called myophrisks. The myophrisks are 
believed to be contractile, and to have the function of changing the volume, and 
hence the density, of the cells, enabling them to sink or float. 

Young cells contain a single nucleus, eccentric in the central capsule; older ones 
have several to many nuclei. 

Haeckel Hsted twenty families in his legion Acantharia. Other authors recognize 
about a half dozen, including the following. 

Family Litholophida Haeckel (1882). Family Astrolophida Haeckel (1887). Spi- 
cules numerous, radiating, not arranged according to Miiller's law. Litholophus, As- 
trolophus, Actinelius, etc. 

Family Chiastolida Haeckel (1887). Spicules ten to twenty, extending clear 
through the body. Chiastolus, Acanthochiasma. 

Family Acanthometrida Haeckel (1862). Acanthometren J. MuUer (1859). Fam- 
ily Acanthonida Haeckel ( 1882). With twenty spicules arranged according to Miiller's 
law; they may be branched, but do not form a continuous network. In most examples, 
as Acanthometron, Xiphacantha, etc., they are equal; in others, as Amphilonche, two 
of the spicules of the equatorial whorl are much longer than the others. 

Family Sphaerocapsida Haeckel (1882). Family Dorataspida Haeckel I.e. Order 
Sphaerophracta Haeckel (1887). Like the foregoing, but the branches of the radiat- 
ing spicules forming a globular network, or two or more concentric networks. Dora- 
taspis, Sphaerocapsa, Lychnaspis. 

Family Diploconida Haeckel (1862). Order Prunophracta Haeckel (1887). Again 
hke the foregoing, but with the eight spicules of the two polar whorls either extended 
or abbreviated. Diploconus, Hexaconus. 

198] The Classification of Lower Organisms 

Order 4. Monopylaria Haeckel in Jenaische Zeitschr. 15: 422 (1882) . 
Order Afowopy/^^n Hertwig Org. Radiolar. 133 (1879). 
Legion Nassellaria with orders Plectellaria and Cyrtellaria, and six suborders, 

Haeckel in Rept. Voy. Challenger Zool. vol. 18 (1887). 
Legion Nassellaria with sublegions Plectellaria and Cyrtellaria and orders Nas- 
soidea, Plectoidea, Stephoidea, and Cyrtoidea, Haeckel Radiolarien H Teil 
Order Monopylida Delage and Herouard Traite Zool. 1: 215 (1896). 
Suborder Nassellaria Doflein. 

Suborder Monopylaria Minchin Protozoa 256 (1912). 
Order Mo7zop};/ga Poche in Arch. Prot. 30: 218 (1913). 
Suborder Monopylea Kudo Handb. Protozool. 261 (1931). 
Suborder Monopylina Hall Protozoology 218 (1953). 
This order is distinguished by a central membrane with one opening, or with a 
single circular field of pores. From this opening or field as a base, there extends into 
the central capsule a large conical body (apparently a bundle of protoplasmic fibers) 
called the porocone. The skeleton varies from none to highly elaborate; it does not 
in any form consist of separate spicules. Its symmetry is dorsiventral, not radial. 
These skeletons are well known as microfossils. 

In this group, under the name of legion Nassellaria, Haeckel placed twenty-six 
families. Other authors recognize about a half dozen. 

Family Nassellida Haeckel (1887). Skeleton none. Cystidium. 
Family Plectonida Haeckel (1887). Family PlectidaYiztcktl (1882), not based 
on a generic name. Skeleton consisting of three arms radiating from a point opposite 
the mouth of the central capsule; sometimes with a fourth forming a caltrop. 

Family Stephanida Haeckel (1887). Family Stephida Haeckel (1882), not based 
on a generic name. Skeleton including a ring in the sagittal plane, often with a basal 
tiipod and with branches and crossbars. Lithocircus, Zygostephanus. This family is 
well represented by microfossils as far back as the Cambrian. 

Family Eucyrtidina Ehrenberg 1847. Family Polycystina Ehrenberg in Abh. 
Akad. Wiss. Berlin ( 1838) : 128 ( 1839), not based on a generic name. Families Hali- 
cryptina and Lithochytridina Ehrenberg 1847. Family Cyrtida Haeckel (1862). 
Order Cyrtoidea, with twelve families, Haeckel (1887). Skeleton a more or less bas- 
ket-shaped network; the root cyrt- in many of the names is Greek KupTr|, a fishing 
basket. Lithocampe, Cryptocalpis, Eucyrtidium, Theoconus, Dictyoconus, and many 
other genera. This group is common as Mesozoic and Cenozoic microfossils, occurring 
mixed with diatoms and silicoflagellates. 

Family Spyridina Ehrenberg 1847. Family Spyrida Haeckel (1882). Order 
Spyroidea, with four families, Haeckel (1887). The skeleton divided by sagittal 
grooves into two lobes. 

Family Cannobotryida Haeckel (1887). Family Botrida Haeckel (1882), not 
based on a generic name. Order Botryoidea, with three families, Haeckel ( 1887) . The 
skeleton divided by three or more longitudinal grooves into as many lobes. 

Order 5. Phaeosphaeria Haeckel in Sitzber. Jenaische Gess. Med. Naturw. 1879: 
Phaeodariae, with orders Phaeocystia, Phaeogromia, Phaeosphaeria. and 
Phaeoconchia Haeckel op. cit. 

Phylum Protoplasta [199 

Order Tripyleen Hertwig Org. Radiolar, 133 (1879). 

Order Phaeodaria Haeckel in Jenaische Zeitschr. 15: 470 (1882). 

Legion Phaeodaria and orders Phaeocystina, Phaeosphaeria, Phaeogromia, and 
Phaeoconchia Haeckel in Rept. Voy. Challenger Zool. vol. 18 (1887). 

Order Tripylea Doflein. 

Suborder Tripylaria Minchin Protozoa 256 (1912). 

Suborder Tripylea Kudo Handb. Protozool. 263 (1931). 

Suborder Tripylina Hall Protozoology 218 (1953). 
In this order, the central capsule is of isobilateral symmetry, having a rather 
small main opening (astropyle) at one end and two smaller openings (parapyles) 
at the other. The openings are located on projections of the central capsule mem- 
brane; inside of each, the protoplasm is so differentiated as to appear to be a conical 
bundle of fibers with the apex at the opening (in contrast to the preceding order, in 
which the base of the conical structure is at the opening) . A mass of variously colored 
bodies, supposedly excreted from the central capsule, lies in the extracapsular cyto- 
plasm about the astroplyle. The skeletons consist in part of organic matter and are 
not well preserved as fossils. 

Borgert (1896, 1900) described nuclear division in Aulacantha. A very large 
number of chromosomes, a matter of several hundred, form a plate which splits into 
two; the two plates move apart in a body of differentiated cytoplasm, but no definite 
spindle, and no centrosomes, were seen. The margins of the plates draw apart faster 
than the middles, with the effect that the plates become saucer-shaped, then bowl- 
shaped, and finally globular, after which nuclear membranes form about them. While 
the nucleus divides, the central capsule membrane becomes constricted by longi- 
tudinal grooves so placed that each daughter central capsule membrane receives 
one parapyle and an astropyle formed from half of the original astropyle. The rudi- 
ments of additional parapyles are first seen as granules in the intracapsular cyto- 
plasm. Each granule grows slightly and becomes "hat-shaped," and migrates so as 
to come into contact with the central capsule membrane at the point appropriate for 
the development of its second parapyle. 

Later, Borgert (1909) described a process in which the nucleus divides repeatedly, 
producing many. The divisions are mitotic, with small numbers of chromosomes, 
perhaps twenty; the eventual products become the nuclei of gametes. There are re- 
ports, in part illustrated with photographs, of similar processes in family Thalassi- 
collida (Hacker, 1907; Huth, 1913). According to Hollande (in Grasse, 1953) the 
small nuclei are those of a parasitic dinoflagellate, Solenodinium. Le Calvez (1935) 
found Coelodendrurn to produce zoospores with a pair of unequal simple flagella. 
They resemble cells of Cryptomonas or of Bodo. 

Haeckel's legion Phaeodaria was of fifteen families. These have been maintained 
by the generality of authors. 

A. Skeleton none or of distinct spicules; cells usually nearly spherical. 
Family Aulacanthida [Aulacanthidae] Haeckel (1879). Aulacantha. 

Family Astracanthida [Astracanthidae] Hacker. Spicules more or less thorny at 
the distal ends. Aulactinium. 

B. Skeleton spherical or of two concentric spheres, with no main opening. 
Family Aulosphaerida Haeckel (1862). Aulosphaera. 

Family Cannosphaerida [Cannosphaeridae] Haeckel (1879). Cromodromys. 
Family Sagosphaerida Haeckel (1887). 

C. Skeleton with a distinct main opening, either nearly spherical, radially sym- 
metrical, or distinctly dorsiventral. 


The Classification of Lower Organisms 

Family Castanellida [Castanellidae] Haeckel (1879). Skeleton nearly globular 
with numerous pores. Castanidium. 

Family Circoporida [Circoporidae] Haeckel (1879). Like the foregoing, but with 
the pores gathered about the bases of radiating spines. Circoporus. 

Family Tuscarorida Haeckel (1887). The main pore on an extended neck, the 
skeleton accordingly flask-shaped. Tuscarora. Tuscarilla. 

D. Shell strongly dorsiventral. 

Family Challengerida [Challengeridae] J. Murray. Shell finely pitted. Chal- 

Family Medusettida Haeckel (1887). Shell smooth or with small spines. Medu- 

E. Shell bilaterally divided into two parts. 
Family Concharida [Concharidae] Haeckel (1879). 

Family Coelodendrida Haeckel (1862). The shell bearing extensive branched 
appendages. Coelodendrum. 

Class 5. SARKODINA (Hertwig and Lesser) Biitschli 

Sarkodina Hertwig and Lesser in Arch. mikr. Anat. 10 Suppl. 43 (1874). 
Class Sarkodina Biitschli in Bronn Kl. u. Ord. Thierreichs 1, 1 Teil: Inhalt ( 1882). 
Class, subclass, etc., Rhizopoda Auctt.; class, subclass, etc. Sarcodina Auctt. 
Amoeboid organisms without flagellate stages, the pseudopodia of the character 
of lobopodia; without skeletons, without or with shells of various materials. 

Fig. 39 — Chaos Protheus: a, b, cells x 25, after the original figures of Pelomyxa 
carolinensis by Wilson (1900); c, mitotic figure x 2,000 after Short (1945). 

Phylum Protoplasta [201 

Hertwig and Lesser took note that the name Rhizopoda was originally applied to 
organisms such as Miliola, which have rhizopodia; they proposed the name Sarko- 
dina for all amoeboid organisms, with Rhizopoda as a subordinate group. Among 
examples of Sarkodina which are not Rhizopoda, they listed first Difflugia, which 
may accordingly be considered the standard genus. 

The Sarkodina as here presented are not assumed to be a natural group. Their 
common characters are probably the outcome of degeneration, by which organisms of 
diverse evolutionary origins have lost their distinctions. 

This assemblage is obviously and superficially divisible into two by the absence or 
presence of shells. The resulting groups are construed as orders. 

Order 1. Nuda Schultze 1854. 

Family Amoebaea Ehrenberg Infusionthierchen 125 (1838). 
Order Lofco^a Carpenter 1861. 

Order Gymnamoebae Haeckel Gen. Morph. 2: xxiv (1866). 
Order AmoebinaKtnt Man. Inf. 1: 27 (1880). 

Suborder Amoebaea Biitschli in Bronn. Kl. u. Ord. Thierreichs 1: 176 (1880). 
Order Gymnamoebida Delage and Herouard (1896). 
Order Chaidea Poche in Arch. Prot. 30: 170 (1913). 

Subclass Amoebaea and order Amoebida Calkins Biol. Prot. 335, 337 (1926). 
Order Amoebaea Kudo Handb. Protozool. 204 (1931). 
Sarkodina without shells. The type is the common amoeba, Amiba diffluens. 
1 Protoplasts not tending to form pseudoplas- 
modial communities. 

2. Free-living Family 1. Amoebaea. 

Family 2. MAYORELLroA. 
Family 3. TnECAMOEBroA. 
Family 4. Hyalodiscida. 

2. Entozoic Family 5. ENDAMOEBroA. 

1. Protoplasts assembling and acting in unison 
in pseudoplasmodial communities. 

2. Parasitic in plants Family 6. Labyrinthulida. 

2. Predatory on bacteria, in air on moist 
surfaces; mostly producing complicated 

fructifications Family 7. Guttulinacea. 

Family 1. Amoebaea Ehrenberg Infusionsthierchen 125 (1838). Family Amoe- 
bidae Bronn 1859. Family Chaidae Poche in Arch. Prot. 30: 171 (1913). Family 
Chaosidae Chatton in Grasse Traite Zool. 1, fasc. 2: 58 (1953). The ordinary free- 
living amoebas. SchaeflFer (1926) limited the family to forms which produce numer- 
ous indefinite granular pseudopodia. There has been much confusion as to the iden- 
tity of the species. There are apparently two species of common large amoebas: 

1. Chaos Protheus L. Syst. Nat. ed. 12: 1326 (1767) {Volvox Chaos L. Syst. Nat. 
ed. 10: 821. 1758. Vibrio Protheus O. F. Muller Verm. Terr, et Fluv. 1: 45. 1773. 
Pelomyxa carolinensis Wilson in American Nat. 34: 535. 1900. Chaos chaos Stiles). 
Schaeffcr identified Pelomyxa carolinensis as the original Chaos Protheus L. It is 
exceptionally large, being macroscopically visible, and is multinucleate. Surely, 
sound nomenclature will apply to this species the name which Linnaeus gave it. 

2. Amiba [Amoeba] diffluens (O. F. Muller) Ehrenberg Infusionsthierchen 127 
(1838) [Proteus diffluens O. F. Muller Animac. Infus. 9. 1786; there is an older genus 

202 ] The Classification of Lower Organisms 

Proteus; Amiba divergens Bory Diet. Class. Hist. Nat. 1: 261. 1822; Amoeba Proteui 
Leidy). It appears that MiiUer intended to rename the Chaos Protheus of Linnaeus; 
that in 1773 he actually did so; but that in 1786 he applied another new name to a 
different organism. Ehrenberg's amended spelling Amoeba, although in general use, 
is not valid as that of a generic name; as Schaeffer suggests, the word may be used as 
a common noun. Amiba diffluens is uninucleate; large, but not visible to the naked 

In nuclear division in the common amoebas the nuclear membrane disappears. 
There are many chromosomes in a blunt-ended spindle. Short (1945) noted a pecu- 
liar twisting of the spindle of Chaos Protheus. 

Schaeffer included in the present family three further genera, Trichamoeba Fro- 
mentel, Polychaos Schaeffer, and Metachaos Schaeffer. Here, in ignorance of its 
relationships, another well-known genus is assigned to this family. 

Pelomyxa, typified by P. palustris Greeff ( 1874) , resembles Chaos Protheus in being 
exceptionally large, macroscopically visible, and multinucleate. It is definitely dif- 
ferent from Chaos Protheus in manner of movement (King and Jahn, 1948) and in 
chemical characters (Andressen and Holter, 1949). 

Minute amoebas moving by means of a single pseudopodium are called Vahlkamp- 
fia. They are believed to have swimming stages with paired equal flagella. If so, they 
do not belong to the present group, but perhaps to the plant kingdom. 

Family 2. Mayorellida [Mayorellidae] Schaeffer in Publ. Carnegie Inst. 345: 47 
(1926). Producing numerous brief conical pseudopodia, but moving by a single large 
clear one. Mayorella, Pontifex, and several other genera proposed by Schaeffer; 
Dactylosphaerium Hertwig and Lesser; Dinamoeba Leidy? The last may be the non- 
flagellate stage of Chactoproteus Stein. 

Family 3. Thecamoebida [Thecamoebidae] Schaeffer op. cit. 83. Amoebas with a 
tough pellicle simulating a shell, moving by the outflow of clear protoplasm at the 
anterior margin. Thccamocba Fromentel. Rugipes Schaeffer. 

Family 4. ''Hyalodiscida [Hyalodiscidae] Poche in Arch. Prot. 30: 182 (1913). 
Family Cochliopodiidae de Saedeleer in Mein. Mus. Roy. Hist. Nat. Belgique 60: 5 
(1934). Similar to the foregoing but without the tough pellicle. Commonly dome- 
shaped, with a row of small pseudopodia projecting from the margin. Hyalodiscus 
and Cochliopodium of Hertwig and Lesser, together with certain genera of Schaeffer. 

Family 5. Endamoebida [Endamoebidae] Calkins. Entozoic amoebas. 

Endamoeba Leidy is found in cockroaches and termites. The nucleus contains no 
karyosome, but many separate granules; in mitosis, definite chromosomes arc formed 
(twelve in E. disparita), but there is apparently no centrosome; at least, no intra- 
desmose is seen (Kirby, 1927). 

Entamoeba Casagrandi and Barbagello, named at nearly the same time as the fore- 
going and regrettably similarly, is widely distributed in invertebrate and vertebrate 
hosts. E. dysenteriae (Councilman and Lafleur) Craig {Endamoeba histolytica Schau- 
dinn) is a serious pathogen to man, the cause of amoebic dysentery. E. coli and E. 
gingivalis are believed to be harmless commensals. The fully mitotic character of 
nuclear division in these organisms was established by Kofoid and Swezy ( 1921, 1922, 
1925). The nucleus contains a small karyosome and an intranuclear centrosome. Mi- 
tosis begins with division of the centrosome into two, which remain connected, as they 
draw apart, by a stainable strand, the intradesmose (the term is of Kofoid and Swczy, 
1921). The karyosome breaks up into chromosomes, six in the species mentioned. 
Spindle fibers connecting these to the centrosomes have been seen; Child (1926) 

Phylum Protoplasta [ 203 

found that the two halves of the spindle swing apart as the centrosomes move apart 
like the legs of a compass being extended. There is no doubt that Endamoeba and 
Entamoeba are generically distinct. 

Endolimax, lodamoeba, and Councilmania occur chiefly in vertebrates and include 
species commensal in man. A refined technique is required to discern the characters 
by which they are distinguished from Entamoeba. Karyamoebina Kofoid and Swezy 
(1924, 1925,) another commensal in man, resembles Vahlkampfia in details of the 
mitotic process, and probably does not belong to the present group. Hydramoeba, 
usually listed in the present family, is not an entozoic organism, but a predator on 

Family 6. Labyrinthulida [Labyrinthuhdae] Haeckel ex Doflein Protozoen 47 
(1901). There is a single genus Labyrinthula Cienkowski, and probably only one 
species, L. macrocystis, parasitic in green and brown algae and in the marine seed 
plant Zostera. The uninucleate cells are spindle-shaped. These cells send out from 
one or both ends fine filaments which writhe in the water. The filaments from differ- 
ent cells coil together and produce "tracks" along which the cells glide. The tracks 
form a network on which the cells may be scattered or gathered into clusters; or the 
cells may abandon their tracks and generate new ones. The nature of the tracks is 
not clear. Possibly they are pseudopodia, on which the cells move by absorbing them 
at one end while generating them at the other. Young (1943) found Labyrinthula 
remarkably indifferent to variations in temperature, reaction, and salinity. 

Family 8. Guttnlinacea [Guttulinaceae] Berlese in Saccardo Sylloge 7: 325 (1888) 
Tribe Dictyosteliaceae Rostafinski Vers. 4 (1873). Sorophoreen with families Gut- 
tulineen and Dictyostcliaceen Zopf Pilzthiere 131-134 (1885). Families Guttulineae 
and Dictyosteliaceae Berlese op. cit. 451. Sappiniaceae Olive in Proc. American Acad. 
37: 334 (1901). Families Sappiniidae, Guttulinidae , and Dictyostelidae Doflein 
1909. Family Acrasidae Poche in Arch. Prot. 30: 177 (1913). Suborder Acrasina 
Hall Protozoology 228 ( 1953 ) . Amoeboid cells predatory on bacteria and other scraps 
of organic matter, in air on moist surfaces, commonly on dung. The cells are capable 
of assembling and moving and going into a resting stage in unison. These organisms 
have generally been included among the Mycetozoa; the resemblance is superficial. 

More recently than Olive, Raper (1940) and Bonner (1944) have surveyed the 
group and studied the behavior. Three families have been maintained, but one appears 
sufiicient to accommodate the seven genera and approximately twenty species. 

Cells of Sappinia are binucleate. They do not necessarily assemble in clusters; a 
single cell may secrete a stalk, by which it is raised into the air, where it rounds up 
and becomes dry. Alternatively, small numbers of cells may assemble and secrete a 
common stalk. The dry cells are "pseudospores" : they are capable of resuming ac- 
tivity without casting off a wall. Hartmann and Nagler (1908) described a peculiar 
sexual process in Sappinia diploidea. 

Guttulina and Guttulinopsis produce larger clusters of resting cells than Sappinia 
does; in Guttulina the resting cells are said to be walled spores. 

Acrasis produces fruits, solitary or clustered, of the form of uniseriate rows of spores 
terminal on stalks consisting of rows of dead cells. 

Distyostelium produces fruits consisting of a column of dead cells bearing a globu- 
lar cluster of spores; Polys phondylium and Coenenia produce slightly more elaborate 
fruits of the same general nature. In Dictyostelium, Raper and Bonner saw that the 
amoeboid active cells, having devoured the available food, gather into a disk-shaped 
mass which may exceed a millimeter in diameter. Wilson (1953) found syngamy, 


The Classification of Lower Organisms 

Fig. 40. — a, Labyrinthula as a parasite in cells of Ectocarpus Mitchelliae x 1,000 
after Karling (1944). b. Cell of Labyrinthula x 2,000 after Young (1943). 
c, d^ Sappinia pedata, active cell and cyst, x 1,000 after Dangeard ( 1896) . e, f, Sap- 
pinia pedata, cluster of pseudospores x 100 and single pseudospore x 1,000 after 
Olive (1902). g, h, Guttulina scssilis, cluster of pseudospores x 100 and individual 
pseudospores x 1,000 after Olive (1902). i-n, Dictyostelium discoideum x 10 after 
Bonner (1944) : i, the pseudoplasmodium; j, it heaps itself up; k, falls toward the 
light and creeps; 1, m, again heaps itself up and becomes a fruit, n. o, p. Fruits of 
Dictyostelium mucoroides; q, of Polysphondylium violaccum; x 10, after Bonner, 
op. cit. 

Phylum Protoplasta [ 205 

karyogamy, and meiosis to occur at this point; the chromosome number (n) is 7. The 
disk changes into a column which bends, and then falls, toward the light, and after- 
ward creeps some distance in the same direction. When this has happened, the fore- 
most cells, being those which were originally in the middle of the disk-shaped mass, 
pile up again to form a sterile stalk perhaps one millimeter tall; the cells behind them 
crawl up the stalk to form the globular mass of spores; the hindmost, being those 
which were last to arrive at the disk-shaped mass, remain behind to form a flange 
about the base of the stalk. 

Order 2. Lampramoebae Haeckel Gen. Morph. 2: xxiv (1866). 
Order Testacea Schultze 1854, non L. (1758). 
Order Thecamoebae Haeckel. 

Order Conchulina Cash and Hopkinson British Freshw. Rhizop. 1: 37 (1905). 
Suborder Testaceolobosa de Saedeleer in Mem. Mus. Roy. Hist. Nat. Belgique 

60: 5 (1934). 
Order Testacida Hall Protozoology 241 (1953). 

Order Testaceolobosa Deflandre in Grasse Traite Zool. 1, fasc. 2: 97 (1953). 
Amoeboid organisms without known flagellate stages, bearing shells and producing 
lobopodia. Various organisms producing rhizopodia or filopodia, traditionally asso- 
ciated with these, have here been placed among Rhizopoda or Heliozoa, as suggested 
by de Saedeleer (1934) and Grasse (1953). Deflandre (in Grasse, op. cit.) distin- 
guishes several families beside the following: 
1. Shell without secreted scales of silica. 

2. Shell of uniform secreted material Family 1. Arcellina. 

2. Shell with imbedded grains of sand Family 2. Difflugiida. 

1. Shell with secreted scales of silica Family 3. NEBELroA. 

Family 1. Arcellina Ehrenberg Infusionsthierchen 129 (1838). Family Arcellidae 
Schultze 1876. Arcella, etc. 

Family 2. Difflugiida [Difflugiidae] Taranek 1881. Difflugia, etc. 
Family 3. Nebelida [Nebelidae] Schouteden 1906. Nebela, the shell beset with 
circular siliceous scales; Quadrula, the scales square; etc. 

Chapter XI 

Phylum 7. FUNGILLI Haeckel 

Order Gregarinae Haeckel Gen. Morph. 2: xxv (1866). 

Class Sporozoa Leuckart Parasiten der Menschen 1, part 1: 241 (1879). 

Phylum FuNGiLLi Haeckel Syst. Phylog. 1: 90 (1894). 

Class Sporozoaria Delage and Herouard Traite Zool. 1 : 254 (1896). 

Subphylum Sporozoa Calkins Biol. Prot. 249 (1926). 

Essentially unicellular organisms (the cells sometimes becoming multinucleate or 
multiple, but remaining undifferentiated except in connection with reproduction); 
commonly with a writhing motion; reproduction usually involving complicated sexual 
processes and the production of walled cysts (spores); flagella absent except some- 
times on the sperms; parasitic in animals. 

The class Sporozoa as originally published by Leuckart included the following 
groups: (a) the gregarines, first described by Dufour (1826) as worms parasitic in 
beetles: the generic name Gregarina Dufour (1828) refers to their occurrence in 
crowds; (b, c) coccidians and psorosperms, different sorts of parasites discovered in 
fishes by J. Miiller and Retzius (1842); and, doubtfully, (d) Miescher's tubes 
{Mieschersche Schlduche), being certain abnormal growths in muscles. The cause of 
the pebrine disease of silkworms, which Nageli (in Caspary, 1857) had named A^o- 
sema Bomhycis, belongs to this group but was not originally included, presumably 
because Nageli had considered it to be a schizomycete. 

It has subsequently become known that almost every species of the animal kingdom 
is parasitized by one or more species of Fungilli. Not all of these parasites, but many, 
are serious pathogens. Thus the Fungilli are a very important group and very num- 
erous. The number of species duly registered by name and description is apparently 
some two or three thousand; this is surely a small fraction of the number which exist. 

The transmission of disease by biting arthropods was first demonstrated when 
Theobald Smith (1893) showed that the Texas fever of cattle, caused by Babesia 
bige?7iina, is transmitted by ticks. 

All who have classified the Sporozoa or Fungilli have recognized two prime sub- 
ordinate groups, the first including the gregarines and coccidians, the second includ- 
ing the organisms which were formerly called psorosperms (Myxosporidia or Neo- 
sporidia). In addition to the main bodies of these groups, there are certain organisms 
which have resisted definite placement and have been assigned sometimes to one of 
the main groups, sometimes to the other, and sometimes to additional main groups. 
In the present work the two main groups are treated as classes and the groups of 
uncertain relationship are included in the first. Clearly, this class is to bear the name 
of Sporozoa Leuckart. Schaudinn's famous paper on parasites in the owl (1903) is 
apparently authority for the widely entertained opinion that this class is artificial, 
representing at least two lines of descent. In fact, the class appears natural with the 
possible exception of some of the poorly known groups. The second class is marked 
by positive specialized characters and is clearly natural; it is not clearly certain that 
the second class is related to the first, and it is accordingly not certain that the phylum 
is natural. The classes are distinguished as follows: 

Phylum Fungilli [ 207 

1. Producing resting cells protected by cell walls 

and not containing polar capsules; or not 

producing resting cells Class 1. Sporozoa. 

1. Producing resting cells whose walls consist 

(at least usually) of modified cells, and 

which contain "polar capsules" enclosing 

coiled threads Class 2. Neosporidia. 

Class 1. SPOROZOA Leuckart 

Class Sporozoa Leuckart Parasiten der Menschen 1, Abt. 1: 241 (1879). 

Subclass Gregarinida Biitschli in Bronn Kl. u. Ord. Thierreichs 1, Abt. 1: Inhalt 

Class Sporozoaria and subclass Rhabdogeniae Delage and Herouard Traite Zool. 
1: 254, 255 (1896). 

Legion Cytosporidia Labbe in Thierreich 5: 3 (1899). 

Subclass Telosporidia Schaudinn in Zool. Jahrb. Anat. 13: 281 (1900). 

Class Telosporidia Calkins Biol. Prot. 421 (1926). 

Class Telosporidca with subclasses Gregarinidia, Coccidia, and Haemosporidia; 
and class Acnidosporidea Hall Protozoology 270, 271, 290, 301, 323 (1953). 

Sous-emhranchement des Sporozoaires, with classes Gregarinomorpha, Coccidio- 
morpha, and Sarcosporidia Grasse Traite Zool. 1, fasc. 2: 545 et seq. (1953). 

Fungilli which produce resting cells protected by cell walls and not containing 
polar capsules; or else do not produce resting cells. 

The nature of the organisms included in this class may be made clear by an 
example, Goussia Schubergi (Schaudinn) comb. nov. [Coccidium Schubergi Schau- 
dinn, 1900). 

Goussia is parasitic in centipedes. Infection is by certain spindle-shaped cells which 
have a certain power of movement, and which make their way to the interior of cells 
of the epithelium of the gut of the host. Each parasitic cell grows and becomes 
globular; it becomes multinucleate; when the host cell dies and breaks up, the para- 
sitic cell divides into many spindle-shaped cells which infect other cells of the gut 

Alternatively, a sexual process takes place. Some of the parasites emerge into the 
gut and do not divide but function as eggs. Others produce numerous cells which 
are more slender than the usual infective cells. These become flagellate, each pro- 
ducing two unequal flagella, and function as sperms. 

The zygote becomes walled. Its nucleus divides twice. Each of the four resulting 
nuclei becomes the nucleus of a walled cyst. The cysts are apparently formed by a 
process of free cell formation: not all of the cytoplasm of the zygote is included in 
them. In each cyst, two of the spindle-shaped infective cells are produced, again ap- 
parently by free cell formation, excluding a part of the cytoplasm. The zygotes, with 
their included cysts and infective cells, pass out with the feces of the host. If a centi- 
pede eats one of them with its food, the infective cells are released to perform their 

No feature of the life cycle described is peculiar to the Sporozoa as contracted 
with other nucleate organisms. Nevertheless, largely by the authority of Schaudinn, 
specialists in Sporozoa use an extensive system of special terms. A familiarity with 
these is necessary to anyone reading about Sporozoa. They include the following: 


The Classification of Lower Organisms 

Sporozoite, the original infective cell. 

Trophozoite, the vegetative individual. 

Nucleogony, the multiplication of nuclei. 

Plasmotomy, multiplication of cells. 

Meront or schizont, the individual in process of dividing to produce further infec- 
tive cells. 

Schizogony or agamogony, the process of dividing to produce infective cells. 

Merozoite or agamete, the infective cell produced from a trophozoite. 

Gamogony, the production of gametes. 

Macrogamete and microgamete mean, of course, egg and sperm; macrogametocyte 
and microgametocyte mean the cells which produce them. 

Sporoblast or sporont, the zygote or other cell inside of which walled cysts are 

Sporogony, the sexual cycle which produces walled cysts. 

Sporulation, the production of walled cysts by asexual processes. 

Spore, the walled cyst. 

Trophozoite (or schizont) and sporont are regarded as the alternating main stages 
in the life cycle of Sporozoa. The point at which meiosis occurs is uncertain. In the 

Fig. 41. — Life cycle of Goussia Schubergi after Schaudinn (1900) : a, sporozoites; 
b-d, developing trophozoites; e, schizogony; f, merozoites; g, young gamctocytes; 
h. i, development of egg; j-m, development of sperms; n, fertilization; o, zygote 
(sporoblast); p-t, development of spores; u, germination of spores. 

Phylum Fungilli [ 209 

monocystid gregarines, Muslow (1911) and Calkins and Bowling (1926) described 
a reduction of the chromosome number immediately before gametogenesis, quite as 
in typical animals. They described reduction as accomplished by a single process of 
nuclear division; to current cytological theory, this is an impossibility. Dobell and 
Jameson (1915), Jameson (1920), and Dobell ( 1925), dealing with organisms of the 
same group and also with the coccodian Aggregata, found meiosis to occur immed- 
iately after karyogamy. They conclude that all nuclei except those of zygotes are hap- 
loid, as among most of the lower plants. 

The coccidian group, to which Goussia belongs, is here treated as primitive among 
Sporozoa because the sperms of this group are flagellate. The detailed structure of 
the flagella is unknown; they appear to resemble those of Bodo and Cryptobia. This 
fact conveys the best available hint as to what may have been the evolutionary origin 
of the Sporozoa. The majority of Sporozoa, having gametes which are alike or 
scarcely differentiated, appear to be derived from forms with markedly differentiated 

The Sporozoa are classified primarily by whether or not the trophozoites are intra- 
cellular; by the occurrence or non-occurrence of asexual reproduction; and by the 
production or non-production of spores in the sense in which the term is used in 
deaHng with this group, that is, of walled cysts. 
1. Sexual reproduction, so far as it is known, 
involving oocytes which produce single large 
eggs and spermatocytes which produce from 
few to many sperms; the organisms multiply- 
ing also asexually. 

2. The gametocytes not attached in pairs. 

3. Producing walled spores Order 1. Oligosporea. 

3. Not producing walled spores. 

4. Intracellular in erythrocytes Order 3. GYMNOSPORiDnoA. 

4. Producing macroscopic bodies 

in muscle Order 4. Dolichocystida. 

2. The gametocytes pairing before gameto- 
genesis; sperms few; with or without 

walled spores Order 2. Polysporea. 

1. Gametes slightly differentiated or undifferen- 
tiated, produced by the gametocytes in more 
or less equal, usually large, numbers. 

2. The organisms multiplying also asex- 

3. Spores producing several sporozoites. . . . Order 5. Schizogregarinida. 
3. Each spore producing one sporozoite. . . . Order 8. Haplosporidhdea. 
2. The organisms not multiplying asexually. 
3. Cells not elongate and divided into 

two parts Order 6. Monogystidea. 

3. Cells elongate and divided into two 

parts Order 7. Polycystidea. 

Order 1. OUgosporea Lankester in Enc. Brit. ed. 9, 19: 855 (1885). 

Tribe Monosporees and groups Disporees and Tetrasporees Schneider in Arch. 
Zool. Exp. Gen. 9: 387 (1881). 

210] The Classification of Loivcr Organisms 

Coccididae, with tribes Monosporea and Oligosporea, Biitschli in Bronn Kl. u. 
Ord. Thierreichs 1: 574, 575 (1882). 

Order Monosporea Lankester op. cit. 854. 

Suborder Coccididae Delage and Herouard Traits Zool. 1 : 278 ( 1896). 

Order Coccidiidia Lahhe in Thierreich 5: 51 (1899). 

Order Coccidiomorpha Doflein Protozoen 95 (1901). 

Order, suborder, or tribe Eimeridea Leger in Arch. Prot. 22 : 80 (1911). 

Order Eimeriidea, suborders Selenococcidinea and Eimeriinea, and tribe Eimer- 
ioidae, Poche in Arch. Prot. 30: 237, 238 (1913). 

Subclass Coccidiomorpha and order Coccidia Calkins Biol. Prot. 435, 436 

Suborder Eimeridea Reichenow in Doflein Lehrb. Prot. ed. 5, 3: 921 (1929). 

Order Eimeriida Hall Protozoology 297 (1953). 
Sporozoa living mostly within epithelial cells of their hosts, multiplying asexually, 
the gametocytes not pairing before gametogenesis, the macrogametocytes producing 
single eggs and the microgametocytes numerous flagellate sperms, the zygotes usually 
producing definite walled spores. 

The organisms of the present order and the following are called coccidians. 
Schneider ( 1881 ) classified them by the number of spores produced in each sporoblast 
(i.e., zygote), either one, two, four, or many. Biitschli and Lankester gave due form 
to Schneider's system. As between their names Monosporea and Oligosporea, the one 
which included the typical example Eimeria is here chosen in preference to the one 
which had page priority. Leger classified these organisms primarily by the number of 
sporozoites per zygote, and distinguished eight families. Here, with the authority, 
for example, of Reichenow (1929) and Kudo (1931), these are reassembled as one 
family to which are appended three others including markedly exceptional or poorly 
known forms. 

Family 1. Eimerida [Eimeridae] Minchin 1903. The typical coccidians. In 
Eimeria Schneider {Coccidium Leuckart) the zygote produces four firmly-walled 
spores each with two sporozoites. The spores are symmetrically ellipsoid and release 
the sporozoites through a terminal pore. Species of this genus parasitize many verte- 
brate hosts, rabbits, sheep, goats, swine, dogs, cats, chickens, turkeys, frogs, and 
fishes. Some of the other genera differ from this as follows: Jarrina, attacking birds, 
is distinguished by spores bearing the pore at the end of a brief neck. Goussia, in cen- 
tipedes, has spores whose walls split lengthwise into two valves. The zygote of Iso- 
spora, in mammals, including man, produces two spores each with four sporozoites; 
that of Caryospora, in snakes, produces one spore with eight sporozoites. Barrouxia, 
in various invertebrates, produces from each zygote numerous bivalved spores each 
containing one sporozoite. 

Family 2. Dobelliida [Dobelliidae] Ikeda. The single known species, Dohcllia bi- 
nuclcata, occurs in a siphuncuHd worm. It exhibits an exception to the characters 
of the order: the male and female gametocytes become attached to each other; the 
male gametocyte, however, produces many sperms, as in the generality of the order. 
Family 3. Aggregatida [Aggrcgatidae] Labbe in Thierreich 5: 6 ( 1899). This fam- 
ily is distinguished by hetcroocism. In Aggrcgata Ebcrthi, vegetative growth and 
multiplication take place in crabs. When these are eaten by squids, the cells either 
develop into single eggs or else divide to produce many sperms. The zygote produces 
about twenty bivalved spores which pass out with the feces and infect crabs. The 
number of sporozoites per spore is variable. There are several other species of Ag- 

Phylum Fungilli [211 

gregata. Various other genera, Merocystis, Hyaloklossia, Myriospora, Caryotropha, 
etc., attacking mussels, polychaet worms, and other marine invertebrates, are as- 
signed to this family although their life cycles are not fully known. 

Family 4. Selenococcidiida [Selenococcidiidae] Poche in Arch. Prot. 30: 238 
(1913) includes the single species Selenococcidium intermedium Leger and Dubosq 
(1910) in the lobster. The vegetative cell is long and slender, and asexual reproduc- 
tion is regularly by transverse division into eight. 

Order 2. Polysporea Lankester in Enc. Brit. ed. 9, 19: 855 (1885). 

Tribe Polysporea Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 576 (1882). 

Suborder Haemosporidae Delage and Herouard Traite Zool. 1 : 284 ( 1896) . 

Order Haemosporidiida Labbe in Thierreich 5: 73 (1899). 

Order, suborder, or tribe Adeleidea Leger in Arch. Prot. 22: 81 (1911). 

Tribe Adeleoidae Poche in Arch. Prot. 30: 239 (1913). 

Order Adeleida with suborders Adeleina and Haemogregarinina Hall Proto- 
zoology 296 (1953). 
It is characteristic of this order that pairs of reproductive cells, essentially mero- 
zoites, which are to become gametocytes, become attached to each other. The mac- 
rogametocyte becomes converted into a single egg; the microgametocyte produces, 
at least usually, four sperms. 

Family 1. Adeleida [Adeleidae] Mesnil in Bull. Inst. Pasteur 1: 480 (1903). 
Chiefly in invertebrates, either in the gut epithelium or in the kidneys, testes, or other 
organs. Zygote usually producing definite spores, these numerous (commonly twenty 
or more), thin-walled, without definite dehiscence mechanism, with two or four 
sporozoites. Adelea and Adelina chiefly in centipedes; Klossia and Orcheobius in 
snails; Klossiella in the kidney of the mouse; Legerella in various arthropods, the zy- 
gote not producing spores but numerous sporozoites. 

Family 2. Haemogregarinida [Haemogregarinidae] Liihe in Mense Handb. Tro- 
penkrankheiten 3: 205 (1906). Heteroecious, with vegetative multiplication in the 
tissues of a vertebrate host. The infection spreads to the erythrocytes of the host, and 
blood-sucking invertebrates are infected by these. Sexual reproduction occurs in the 
invertebrate host. Production of spores is suppressed; the zygote produces numerous 
sporozoites. Haemogregarina Danilewski (1885; Drepanidium Lankester 1882, non 
Ehrenberg 1861) in turtles, frogs, fishes, transmitted by leeches; Hepatozoon in ro- 
dents, Karyolysus in lizards, transmitted by mites. 

Order 3. Gymnosporidiida Labbe in Thierreich 5: 77 (1899). 

Suborder Gymnosporidae Delage and Herouard Traite Zool. 1: 284 (1896). 
Suborder Haemosporidia Doflein Protozoen 121 (1901). 
Order Haemosporidia Calkins Biol. Prot. 441 (1926). 

Subclass Haemosporidia with orders Plasmodiida and Babesiida Hall Proto- 
zoology 301, 302, 306 (1953). 
In this order the vegetative cells occur in vertebrates and infect the erythrocytes. 
Sexual reproduction, so far as it has been discovered, occurs in blood-sucking arthro- 
pods. The gametocytes do not become associated in pairs; the male gametocytes pro- 
duce numerous spirochaet-like sperms by a process of budding. In the zygote, the 
nucleus undergoes a series of divisions, after which numerous naked uninucleate 
sporozoites are budded off from the surface. There are no walled spores. 


The Classification of Lower Organisms 

The name Haemosporidia, commonly applied to this order, appears to belong by 
priority to the preceding. 

Schaudinn (1903) was disposed to connect this order with the trypanosomes, while 
connecting the coccidians with Bodo and Cryptobia. This view has been entertained 
by Liihe (inMense, 1906), Woodcock (1909), and Leger (1910). In spite of authority 
thus good, it appears far-fetched. The Gymnosporidiida are of the same general na- 
ture as the Aggregatida, Adeleida, and Haemogregarinida. 

Fig. 42. — a-m. Life cycle of Plasmodium compiled from various sources: a, infec- 
tion of an erythrocyte by a sporozoite; b-e, trophozoites, plasmotomy, and mero- 
zoites; f, spermatocyte; g, oocyte; h, production of sperms; i, fertilization; j, k, pro- 
duction of sporozoites in cells of the gut epithelium of the mosquito; 1, sporozoites; 
m sporozoites entering the salivary gland of the mosquito, n-q. Stages of division 
of cells of Babesia bigemina in erythrocytes of cattle x 2,000 after Dennis (1930). 

The Gymnosporidiida are organized, somewhat arbitrarily, as three families. 

Family 1. Halteridiida [Halteridiidae]Hartmann and Jollos 1910. Family Leu- 
cocytozoidae Hartmann and Jollos. Family Hacmoproteidae Doflcin. Hacmoproteus 
Kruse {Haltcridium Labbc) occurs in reptiles and birds. Vegetative growth and re- 
production occur in tissue cells. Some of the merozoites infect erythrocytes, and are 
believed to become gametocytes, and to develop no further unless swallowed by some 
blood-sucking arthropod. In the best known example, H. Columbae of pigeons 

Phylum Fungilli [ 213 

(Argao, 1908), the alternate host is a fly. In the gut of the fly, the spermatocytes pro- 
duce the elongate sperms as outgrowths. The zygotes make their way into the wall of 
the gut of the fly, grow, and produce very numerous sporozoites. These migrate to the 
salivary gland, from which they are injected into pigeons. 

Leucocytozoon attacks birds; its cells become fairly large in certain blood cells 
which become colorless and spindle-shaped. 

Family 2. Plasmodida [Plasmodidae] Mesnil in Bull. Inst. Pasteur 1: 480 (1903). 
The malaria organisms, differing from Haemoproteus in that they multiply in the 
erythrocytes of their hosts. With a few obscure exceptions, the species are construed 
as a single genus Plasmodium. Three species attack man; they have perhaps done 
mankind more injury than any comparable group of living creatures. Several com- 
paratively poorly known species attack apes and monkeys. The alternate hosts of all 
species are mosquitoes of the genus Anopheles. 

The vegetative individuals complete their growth within erythrocytes of their hosts 
in more or less definite periods of time, and undergo multiple division; the erythro- 
cytes then break up and release the merozoites. The chill and fever of malaria are as- 
sociated with the destruction of erythrocytes. In the ordinary form of malaria, called 
tertian malaria, development requires forty-eight hours, and the chill and fever occur 
every other day. Another form, called malignant tertian or tropical malaria, exhibits 
the same rhythm; it is distinguished by details of the appearance of the infected 
erythrocytes. In the third form of malaria in man, called quartan, development re- 
quires 72 hours, and the chill and fever occur every third day. 

The course of development in the mosquito is quite like that of Haemoproteus 
Columbae in the fly. Some of the parasites inside the erythrocytes are gametocytes; 
each female gametocyte in an erythrocyte swallowed by a mosquito develops into a 
single egg, while each male gametocyte buds off several spirochaet-like sperms. The 
fertilized eggs are able to move. They break into the epithelium of the gut of the 
mosquito, grow into large globes, and become multinucleate; their protoplasts divide 
into numerous masses of protoplasm each of which buds off large numbers of sporozo- 
ites. The sporozoites are released into the body cavity of the mosquito, migrate to 
the salivary gland, and are injected into whatever animal the mosquito may bite. 

The scientific names usually applied to the three species which cause human 
malaria are not valid by priority. Extensive synonymy is given by Sabrosky and 
Usinger, in their application to the International Commission on Zoological Nomen- 
clature for action arbitrarily maintaining the current names (1944), and in the 
report by Hemming (1950) of the action of the Commission. 

Certain structures in the erythrocytes of malaria patients were first recognized as 
parasites by Laveran, 1880, who, in 1881, named them Oscillaria malariae. The 
organism is believed to have been that of malignant tertian or tropical malaria. The 
word Plasmodium, properly designating a certain type of body, was applied by Mar- 
chiafava and Celli 1885, in the combination Plasmodium malariae, believed also 
originally to have designated the agent of malignant tertian malaria. Feletti and 
Grassi, 1889, introduced the generic name Haemamoeba, with two species, H. vi- 
vax, the agent of tertian malaria, and H. malariae, that of quartan malaria; it is be- 
lieved that the latter epithet was applied under the misapprehension that this was the 
organism which Marchiafava and Celli had named. It appears that Liihe, 1900, is 
responsible for the currently used names: 

Plasmodium vivax, the organism of tertian malaria; 

P. malariae, that of quartan malaria; 

214] The Classification of Lower Organisms 

P. falciparum, that of malignant tertian. 

In order that a great mass of literature may be read without confusion, it is ex- 
pedient that these names be arbitrarily maintained. The International Commission 
of Zoological Nomenclature has duly taken action to this effect. 

Family 3. Babesiida [Babesiidae] Poche in Arch. Prot. 30: 241 (1913). Family 
Theileridae du Toit in Arch. Prot. 39: 94 (1918). Minute intracellular parasites 
transmitted by arthropods; sexual reproduction unknown. Theileria Bettencourt et al. 
causes a fever of cattle in Africa; the parasites multiply in the tissue cells and spread 
to the ery'throcytes, by which ticks are infected. Babesia Stercovici [Piro plasma Pat- 
ton) is similar, but the parasites multiply in the erythrocytes. B. bigemina causes 
the Texas fever of cattle. 

The minute nucleus of Babesia bigemina is largely filled by a single granule, a 
karyosome. This is connected by a rhizoplast to an extranuclear granule which has 
been identified as a blepharoplast, although no flagellum is present. In nuclear divi- 
sion, as described by Dennis (1930), the blepharoplast divides; the rhizoplast splits; 
the nucleus widens, the karyosome becoming a rod; karyosome, nucleus, and cell 
undergo constriction. No chromosomes are seen. 

If Bartonella bacilliformis, the agent of the disease variously known as verruga 
peruana, Oroya fever, or Carrion's disease, is not a bacterium, perhaps it may be 
placed in or near this family. 

Order 4. Dolichocystida Delage and Herouard Traite Zool. 1 : 289 (1896). 
Sarcosporidia Balbiani 1882. 
Class Sarcosporidia Biitschli in Bronn Kl. u. Ord. Thierreichs 1, Abt. 1 : Inhalt 

Subclass Sarcocystidca Lankester in Enc. Brit. ed. 9, 19: 855 (1885). 
Order Sarcosporidia Doflein Protozoen 214 (1901). 
Order Sarcocystidca Poche in Arch. Prot. 30: 245 (1913). 
Subclass Sarcosporidia Calkins Biol. Prot. 461 (1926). 
The characters are those of the single family and genus: 
Family Sarcocystida [Sarcocystidae] Poche in Arch. Prot. 30: 245 (1913). 
Sarcocystis Lankester produces the Mieschersche Schlauche, macroscopically visible 
bodies, globular, fusiform, or filiform, of dimensions up to several millimeters, in 
muscles of animals. The several supposed species, from mice, sheep, swine, deer, etc., 
are not morphologically distinguishable. Miescher observed these things in mice, in 
which they are called Sarcocystis Muris; material from swine is called S. Miescher- 

The visible body is a mass of cells, the whole walled by modified muscle of the 
host. The mass originates as a single cell which divides repeatedly; the ultimate 
division products are crescent-shaped uninucleate reproductive cells. Erdmann (1910) 
observed the infection of epithelial cells of the gut of mice. Each infective cell grew 
and divided into several, v\'hich made their way, or were carried, to the muscles, where 
they gave rise to the Mieschersche Schlauche. Crawley (1914, 1916), on the other 
hand, found the infective cells to be gametocytes. In cells of the gut epithelium of 
the host, they may be converted as wholes into eggs, or else may give rise to numerous 
elongate sperms. These conflicting observations could be explained by an alternation 
of sexual and asexual generations, but the point is not established. 

Phylum Fungilli [ 215 

Order 5. Schizogregarinida Calkins Biol. Prot. 433 (1926). 
Amoebosporidies Schneider. 

Amoebosporidia Labbe in Thierreich 5: 120 (1899). 
Suborder ^mo^feo^/^oncfm Doflein Protozoan 171 (1901). 
^iuhordev Schizocystinea Poche in Arch. Prot. 30: 233 (1913). 
Suborder Schizogregarinaria Reichenow in Doflein Lehrb. Prot. ed. 5, 3 : 872 

Orders Archegregarina and Neogregarina Grasse Traite Zool. 1, fasc. 2: 622, 
665 (1953). 
The Sporozoa previously considered, particularly those of the first two orders, are 
called coccidians; those of the present order and the two which follow are called 
gregarines. The latter are characterized (not without exceptions) by inter- instead 
of intra-cellular active stages, and by the production of numerous gametes, alike or 
not strongly differentiated, from paired scarcely differentiated gametocytes. The 
present order includes the gregarines which exhibit asexual reproduction. They are 
a rather miscellaneous assemblage. 

Family 1. Schizocystida [Schizocystidae] Leger and Duboscq in Arch. Prot. 12: 
102 (1908). Family Monoschizae V^eiser in ]our. Protozool. 2: 10 (1955), including 
the two following families. In marine worms and other invertebrates. The sporozoites 
enlarge in the host and become multinucleate individuals which reproduce freely by 
producing uninucleate buds. Some of these buds continue the infection directly; 
others become attached in pairs, each pair secreting a common cyst wall. Each of the 
individuals in the cyst become multinucleate and buds off numerous uninucleate 
gametes. The zygotes become walled spores which are cast out with the feces of the 
host, to infect others which ingest them. Each produces eight sporozoites. Schizo- 
cystis, Siedleckia. 

Family 2. Seleniida [Seleniidae] Brasil in Arch. Prot. 8: 394 (1907). In marine 
worms. Vegetative individuals notably long and slender; spores spiny, with four 
sporozoites. Selenidiu7n, Meroselenidium. 

Family 3. Merogregarinida [Merogregarinidae] Fantham 1908. Family Caul- 
leryellidae Keilin. Merogregarina, Caulleryella, Tipulocystis. 

Family 4. Spirocystida [Spirocystidae] Calkins Biol. Prot. 435 (1925). Family 
Spirocystidees Leger and Duboscq in Arch. Prot. 35: 210 (1915). In earthworms. 
Spores containing a solitary sporozoite which escapes through a pore. Spirocystis. 

Family 5. Ophryocystida [Ophryocystidae] Leger and Duboscq in Arch. Prot. 12: 
102 (1908). Family Amoebosporidiidae Brasil (1907), not based on a generic name. 
Family Dischizac Weiser in Jour. Protozool 2: 10 (1955). In Ophryocystis Schneider 
(Leger, 1907), the vegetative individuals, attached to the walls of the Malpighian 
tubules of beetles, grow and become multinucleate and send out branches whose ends 
develop into additional individuals. Eventually, different individuals become at- 
tached in pairs. Each of these individuals buds off a single uninucleate gamete. The 
remaining protoplasm of the gametocytes forms a protective sheath around the zygote, 
which becomes a single spore with eight sporozoites. 

Order 6. Monocystidea Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 574 (1882). 

Order Haplocyta Lankester in Enc. Brit. ed. 9, 19: 853 (1885). 

Suborder Acephalina Labbe in Thierreich 5 : 37 ( 1899) . 
Organisms of the character of gregarines, not multiplying asexually, the vegetative 
individuals not elongate and divided into serial parts. 

216] The Classification of Lower Organisms 

The genus which is best known is Monocystis Stein, including several species which 
are common in earthworms. The cells grow within epithelial cells of the seminal fun- 
nels; they and their nuclei reach considerable sizes without dividing. At maturity, 
they escape into the seminal vescicles, where they form pairs, each pair secreting a 
common cyst wall. The pairing and encystment were observed, more definitely of the 
related genus Zygocystis than of Monocystis, by Stein (1848). The nuclei of the 
paired cells divide. Several observers, as Brasil (1905) and Mulsow (1911); also, as 
to related genera, Jameson (1920) and Noble (1938); have observed peculiarities in 
the first nuclear division. The peculiarities amount to this, that the large nucleus 
breaks up and, for the most part, undergoes dissolution, leaving a small number of 
definite chromosomes to undergo normal mitosis in a spindle. Repeated subsequent 
divisions are of normal character. The numerous nuclei thus produced become those 
of gametes which are budded oflF from the surfaces of the gametocytes. This was first 
observed by Wolters (1891). The gametes from the respective paired cells are pre- 
sumably always of different mating types, and are usually visibly differentiated, 
larger and smaller. Each zygote becomes a spindle-shaped walled spore; the enucleate 
remainder of the gametocytes provides nourishment during their development. Each 
spore produces eight sporozoites. 

The number of known species of Monocystidea is of the order of 150. The majority 
occur in annelid worms; others attack flatworms, echinoderms, insects, tunicates, and 
other invertebrates. Bhatia (1930) distinguished twelve famiHes which are here 
merely listed. 

A. The two ends of the spore alike. 

Family 1. Monocystida [Monocystidae] (Biitschli) Poche in Arch. Prot. 30: 236 
(1913). Family Monocystiden Stein in Arch. Anat. Phys. 1848: 187 (1848). Mono- 
cystidae Biitschli ( 1882). Monocystis, etc. 

Family 2. Rhynchocystida [Rhynchocystidae] Bhatia in Parasitology 22: 158 
(1930). Rhynchocystis. 

Family 3. Stomatophorida [Stomatophoridae] Bhatia op. cit. 159. Stomatophora, 
Choanocystis, etc. 

Family 4. Zygocystida [Zygocystidae] Bhatia op. cit. 160. Zygocystis, Pleurocystis. 

Family 5. Akinetocystida [Akinetocystidae] Bhatia op. cit. 160. Akinetocystis. 

Family 6. Syncystida [Syncystidae] Bhatia op. cit. 161. Syncystis. 

Family 7. Diplocystida [Diplocystidae] Bhatia op. cit. 161. Diplocystis, Lankcsteria. 

Family 8. Schaudinellida [Schaudinellidae] Poche in Arch. Prot.' 30: 236 (1913). 

B. The ends of the spores differentiated. 

Family 9. Doliocystida [Doliocystidae] Labbe in Thierreich 5: 33 (1899). Family 
Lecudinidae Kamm. Lxcndina Mingazzini {Doliocystis Legcr). 

Family 10. Urosporida [Urosporidae] Woodcock 1906. Family Choanosporidae 
Dogiel. Gonospora; Lithocystis; Urospora, the spores with long tails; Ceratospora; 
Pterospora, the spores with longitudinal flanges. 

Family 11. Ganymedida [Ganymcdidae] J. S. Iluxlcy in Quart. Jour. Micr. Sci. 
n..s. 55: 169 (1910). Ganymcdcs. 

Family 12. Allantocystidae [Allantocystidae] Bhatia op. cit. 163. Allantocystis. 

Order 7. Polycystidea Biitschli in Bronn Kl. u. Ord. Thicrreichs 1: 578 (1882). 
Order Grcgarinae Haeckel Gen. Morph. 2: xxv (1866), the mere plural of a 
generic name. 

Phylum Fungilli [217 

Subclass Gregarinida Biitschli op. cit. Inhalt (1882). 

Order Septata Lankester in Enc. Brit. ed. 9, 19: 853 (1885). 

Order Brachycystida, suborder Gregarinidae, and tribe Cephalina or Folycystina 
Delage and Herouard Traite Zool. 1: 255, 256, 269 (1896). 

Order Gregarinida Labbe in Thierreich 5: 4 (1899). 

^uhordtr Eugregarinaria Doflein Protozoen 160 (1901). 

Order Gregarinoidca Minchin (1912). 

Suborder Gregarininea and tribe Gregarinoidae Poche in Arch. Prot. 30: 234 

Subclass Gregarinida, order Eugregarinida, and suborder Cephalina Calkins Biol. 
Prot. 422, 428 (1926). 
The typical gregarines, the vegetative cells elongate and divided by more or less 
definite constrictions into two (or, occasionally, more than two) parts; not repro- 
ducing asexually. 

Typical gregarines occur chiefly in insects. The vegetative cell consists of an an- 
terior portion (protomerite) serving for attachment and a posterior portion (deuto- 
merite), containing the nucleus, lying in the gut cavity of the host. Both parts have 
a thick outer layer, commonly differentiated upon the protomerite into a more or 
less elaborate knob, the epimerite. Longitudinal fibrils, presumably contractile, are 
present. The cells writhe actively. 

The individuals are commonly found in pairs, one member attached to the epi- 
thelium of the gut, the other to the posterior end of the first. This arrangement is 
produced by active self-placement on the part of the second member. When both are 
mature, they take common action to produce a globular cyst. The protoplasts remain 
distinct until both have become multinucleate, after which they produce numerous 
gametes. In some forms, as Nina, studied by Goodrich (1938), all of the gametes 
migrate from one cell, recognizably male, into the other, the female cell; the male 
cell is left empty and is compressed or crushed by the growth of the zygotes in the 
female cell. The zygotes are spores, usually fusiform, and usually producing sporo- 
zoites by eights. In Gregarina and Gamocystis, an inner layer of the cyst wall is so 
modelled as to form tubes (sporoducts) running from the surface to the interior. 
When the spores are ripe, the sporoducts become extroverted and the spores are ex- 
truded through them in uniseriate rows. In connection with this behavior, the spores 
have flat ends like barrels. 

Family 1. Stenophorida [Stenophoridae] Crawley 1903. Protomerite a mere 
knob. Stenophora. 

Family 2. Gregarinida [Gregarinidae] Greene 1859. Family Gregarinarien Stein 
in Arch. Anat. Phys. 1948: 187 (1848). Gregarines which are without epimerites 
and are not notably elongate. There are about a dozen genera. Cysts without sporo- 
ducts: Hirmocystis, Hyalospora, Cnemidospora. Cysts with sporoducts, the spores 
barrel-shaped: Gregarina, Gamocystis. The earliest observations of Sporozoa were by 
Dufour ( 1826) , who, studying the anatomy of insects, found them in the gut of beetles. 
He took them for worms and illustrated an individual with an epimerite, which he 
took for a sucker. Later (1828) he applied names, Gregarina conica to the form first 
seen, G. ovata to a form without an epimerite found in the forficule, i.e., in an ortho- 
pteran. The former does not belong to the genus Gregarina as subsequently construed; 
it appears to be a member of the family Actinocephalida. Gregarina ovata should be 
regarded as the type of Gregarina, but the genus has usually been interpreted by G. 
cuneata, which Stein observed in cockroaches. 

218] The Classification of Lower Organisms 

Family 3. Didymophyida [Didymophyidae] Wasilewski 1896. Family Didymo- 
phyiden Stein (1848). Like the foregoing, but the cells extremely elongate. Didymo- 

Family 4. Acanthosporida [Acanthosporidae] Labbe in Thierreich 5: 27 (1899). 
The spores with polar or equatorial bristles. Acanthospora. 

Family 5. Stylocephalida [Stylocephalidae] Ellis 1912. Family Stylorhynchidae 
Labbe op. cit. 30, based on a generic name which is a later homonym. Epimerite 
elongate with a small terminal knob. Stylocephalus. 

Family 6. Actinocephalida [Actinocephalidae] Wasilewski 1896. Epimerite with 
thorns. Numerous genera, Sciadophora, Acanthorhynchus, Actinocephalus, Hoplo- 
rhynchus, Pileoccphalus, etc. 

Family 7. Menosporida [Menosporidae] Labbe op. cit. 29. Epimerite with a long 
stalk, distally branched and bearing appendages. Menospora. 

Family 8. Dactylophorida [Dactylophoridae] Wasilewski 1896. Epimerite dis- 
tally broadened, clinging to the host epithelium by means of numerous filiform pro- 
cesses. Dactylophorus, Nina [Pterocephalus), etc. 

Family 9. Porosporida [Porosporidae] Labbe op. cit. 7. Heteroecious: in Porospora, 
the gregarinoid stage occurs in crabs and the production of spores occurs in mussels. 
The spores contain a single sporozoite and open through a pore. 

Order 8. Haplosporidiidea Poche in Arch. Prot. 30: 178 (1913). 
Order Aplosporidies Caullery and Mesnil 1899. 
Order H aplosporidies Caullery and Mesnil in Arch. Zool. Exp. Gen. ser. 4, 4: 

104 (1905). 
Order Haplosporidia Auctt., the mere plural of a generic name. 
Subclass Haplosporidia Hall Protozoology 326 (1953). 
Unicellular intracellular parasites, the cells becoming multinucleate and multiply- 
ing by fragmentation, producing walled spores which germinate by releasing the 
protoplasts as single sporozoites. 

The vegetative body is of the type properly called a plasmodium. The nuclei and 
the process of division, described by Granata (1914) are characteristic. The resting 
nucleus contains an "axial rod" as well as a nucleolus-like body. In mitosis the axial 
rod becomes converted into an intranuclear spindle. Individual chromosomes have 
not been seen; the chromatin gathers in a mass about the middle of the spindle (the 
figures are curiously diatom-like). The mass of chromatin, the nucleolus-like body, 
and the entire nucleus, divide by constriction; the ends of the spindle persist as the 
axial rods of the daughter nuclei. Eventually, the plasmodium secretes a thin wall 
and the protoplast divides into uninucleate naked cells. Granata found that these 
cells are gametes, and that conjugation takes place among gametes produced by the 
same plasmodium. The zygotes become walled spores which germinate by casting 
off a circular operculum and releasing the contents. If the life cycle is correctly 
understood, we may suppose that these organisms are degenerate gregarinos. 

In the present state of knowledge, it will be as well to treat the typical haplo- 
sporidians as a single family: 

Family Haplosporidiida [Haplosporidiidae] Caullery and Mesnil in Arch. Zool. 
Exp. Gen. ser. 4, 4: 106 (1905). Families Bartramiidae and Coelosporidiidae Caul- 
lery and Mesnil op. cit. 107. Characters of the order. Haplosporidium (spores with 
appendages at both ends) and Urosporidium (spores with a single appendage) attack 

Phylum Fungilli [219 

chiefly annelid worms. Bartramia attacks rotifers; Ichthyosporidium is a serious 
parasite of fishes; Coelosporidium attacks cockroaches. 

The following family, of uncertain position, may tentatively be associated with the 
Haplosporidiidea : 

Family Metchnikovellida [Metchnikovellidae] Caullery and Mesnil in Compt. 
Rend. Soc. Biol. 77: 527 (1914), Ann. Inst. Pasteur 33: 214 (1919). Secondary 
parasites, intracellular in gregarines; cells naked at first, with very minute nuclei, 
which become numerous, later converted into walled cysts of characteristic form, the 
protoplasts undergoing division into uninucleate infective cells. Mctchnikovella, 
Amphiamblys, Amphiacantha. 

Class 2. NEOSPOR!D!A (Schaudinn) Calkins 

Myxosporidia Biitschli in Zool. Jahresber. 1880: 162 (1881). 

Subclass Myxosporidia Biitschli in Bronn Kl. u. Ord. Thierreichs 1, Abt. 1 : Inhalt 

Subclass Amoebogeniae Delage and Herouard Traite Zool. 1: 291 (1896). 

Subclass Neosporidia Schaudinn in Zool. Jahrb. Anat. 13: 281 (1900). 

Order Cnidosporidia Doflein Protozoen 177 (1901). 

Class Cnidosporidia Poche in Arch. Prot. 30: 224 (1913). 

Class Neosporidia and subclass Cnidosporidia Calkins Biol. Prot. 445, 448 (1926). 

Subphylum Cnidosporidia Grasse Traite Zool. 1, fasc. 1: 129 (1952). 

Class Cnidosporidea Hall Protozoology 311 (1953). 

Fungilli whose resting cells contain polar capsules; are walled, at least usually, 
by a layer of modified cells; and, in most examples, release a single infective cell. 

As a general rule, the vegetative bodies of Neosporidia are plasmodia, i.e., naked 
multinucleate bodies, usually freely capable of asexual reproduction by internal or 
external budding. An entire small plasmodium may become converted into one or two 
spores, or the spores may be cut out internally and produced continually. The spores, 
at least in the two better-known orders, are structures formed from several cells; they 
are not homologous with the spores of the proper Sporozoa. In most examples, only 
one of the cells involved in the formation of a spore is fertile, and only one infective 
protoplast is released on germination. Of the sterile cells, one or more become con- 
verted into the structures called polar capsules. These resemble the nematocysts of 
coelenterates : they contain a coiled hollow thread capable of swift extroversion. 
Extroversion occurs during germination. Its significance is unknown. The presence 
of polar capsules marks the class as a natural group. 

Three orders are recognized: 

1. Spores covered by two valves formed from 

accessory cells Order 1. Phaenocystes. 

1. Spores covered by three valves formed from 

accessory cells Order 2. Actinomyxida, 

1. Spores very minute, with a continuous mem- 
brane Order 3. Cryptocystes. 

Order 1. Phaenocytes Gurley in Bull. U. S. Fish Comm. 11 : 410 (1893). 
Order N^maiocj^^ffrfa Delage and Herouard Traite Zool. 1: 291 (1896). 
Order Phaenocystida Labbe in Thierreich 5 : 85 (1899). 


The Classification of Lower Organisms 

Order Cnidosporidia Doflein Protozoen 177 (1901). 
Order Myxosporidia Calkins Biol. Prot. 449 (1926). 
Most species of this order parasitize fishes, living either in internal cavities or in 
the tissue cells; fewer than a dozen species are known from miscellaneous other ani- 
mals, amphibia, reptiles, insects, and worms. Most of these parasites are not 
extremely injurious. 

The infective protoplast which issues from a spore is, at least usually, binucleate. 
The nuclei fuse and the fusion nucleus divides repeatedly as the plasmodium grows. 

Fig. 43.— Diagram of the life cycle of Myxoceros Blennius after E. Noble (1941). 

In the examples which are believed to be more primitive, the plasmodia are freely 
capable of budding, and the mature plasmodia are rather small and are converted as 
wholes into single or paired spores. In the remaining examples, the plasmodia do not 
multiply by budding, but produce spores continually. 

Noble (1941) described the mitotic process. There is a rather large intranuclear 
centrosome, which divides, the daughter centrosomes moving to opposite sides of 
the nuclear cavity. Four chromosomes appear; this is apparently constant throughout 
the order. The nuclear membrane and the centrosomes disappear. No spindle has 
been seen. The chromosomes divide, and the daughter chromosomes move apart and 
melt into two masses. The masses swell, a nuclear membrane appears about each, 
and a centrosome appears inside of each. 

Phylum Fungilli [221 

The spore-forming structure (sporoblast) is a protoplast with several nuclei; it is 
either a whole small plasmodium, or half of one, or a protoplast cut out endogen- 
ously within a plasmodium. Two of the nuclei are set apart in cells which become 
converted into the valves of the spore. Two or four are set apart in cells which become 
converted into polar capsules. Two, of which it is established that they have two 
chromosomes each, are the nuclei of the infective protoplast. 

In a review of the literature as to Hfe cycles, Noble (1944) remarks as follows. "A 
survey of the literature reveals that there is little agreement on the details of nuclear 
changes in the Myxosporidia. Some authors maintain that the cycle is mainly haploid, 
others have described a diploid cycle. Some reports indicate that there are two reduc- 
tion divisions and two zygotes in one cycle. When only one zygote is reported the 
reduction division in one case occurs just before fertilization, in another case it occurs 
just after fertilization. Some authors have maintained that there is no sexual process." 
Noble's own conclusions include the following. The organisms are diploid at most 
stages. The meiotic divisions are among those by which the sporoblast becomes multi- 
nucleate. The two haploid nuclei of the spore, which unite after germination, are 
derived from a single diploid nucleus. Authors who have described fusions of proto- 
plasts, or transfers of nuclei from one protoplast to another, have had no evidence 
beyond an understandable unwillingness to accept fusions of sister nuclei. 

Nearly two hundred species of the present order are listed in the monograph of 
Kudo (1920), who established three suborders. 

A. Valves conical, spores biconic (suborder Eurysporea Kudo). 

Family 1. Myxoceratida nom. nov. Family Ceratomyxidae Doflein Protozoen 182 
(1901), based on a generic name which is a later homonym. Characters of the sub- 
order. Myxoceros nom. nov. [Ccratomyxa Thelohan 1892, non Ceratiomyxa Schroter 
1889; if ever names are homonymous without being absolutely identical, these are.) 
Some thirty-five species; the type is M. sphaerulosa (Thelohan) comb, nov.; Noble 
studied mitosis in M. Blennius (Noble) comb. nov. Leptotheca, Myxoproteus, War- 
dia, Mitraspora. 

B. Valves hemispherical, spores spherical (suborder Sphaerosporea Kudo) . 
Family 2. Chloromyxida [Chloromyxidae] Gurley in Bull. U. S. Fish Comm. 1 1 : 

418 (1893). Chloromyxees Thelohan in Bull. Soc. Philomath. Paris ser. 8, 4: 176 
(1892). Chloromyxea Braun in Centralbl. Bakt. 14: 739 (1893). With four polar 
capsules. Chloromyxum. 

Family S.Sphaerosporida [Sphaerosporidae] Davis 1917. With two polar cap- 
sules. Sphaerospora, Sinuolinea. 

C. Valves saucer- or boat-shaped, spores disk-shaped or fusiform (suborder 
Platysporea Kudo ) . 

Family 4. Myxidiida [Myxidiidae] Gurley op. cit. 420. Myxidiees Thelohan op. 
cit. 175. Myxidiea Braun I.e. Myxidium, Sphaeromyxa, Zschokkella. 

Family 5. Coccomyxida [Coccomyxidae] Leger and Hesse 1907. Coccomyxa. 

Family 6. Myxosomatida [Myxosomatidae] Poche in Arch. Prot. 30: 230 (1913). 
Myxosoma, Lentospora. 

Family 7. Myxobolida [Myxobolidae] Gurley op. cit. 413. Myxobolees Thelohan 
op. cit. 176. Myxobolea Braun I.e. Myxoboliis, Henneguya, Hoferellus. 

Order 2. Actinomyxida Stole 1911. 

This order includes about a dozen parasites in annelid worms. A plasmodial stage 
and asexual reproduction are believed not to occur; the infective protoplast grows 

222 ] The Classification of Lower Organisms 

into an individual whose one or two nuclei remain undivided until the commence- 
ment of the ill-understood process by which the complicated spores, with three valves 
and three polar capsules, are produced. 

Family 1. Tetractinomyxida [Tetractinomyxidae] Poche in Arch. Prot. 30: 231 
(1913). Family Haploactinomyxidae Granata in Arch. Prot. 50: 205 (1925). Spores 
subglobular, with a single binucleate sporozoite. Tetractinomyxon. 

Family 2. Synactinomyxida [Synactinomyxidae] Poche I.e. Family Euactinomyxi- 
dae Granata I.e. Family Triactinomyxidae Kudo Handb. Protozool. 314 (1931). 
Spores producing eight or more sporozoites. S phaeractinomyxon and Neactinomyxon, 
the spores subglobular. Synactinomyxon, with two of the valves protruding as con- 
siderable horns, the whole horse-shoe shaped. Triactinomyxon and Hexactinomyxon, 
all three valves drawn out into long horns, the whole caltrop- or anchor-shaped. 

Order 3. Cryptocystes Gurley in Bull. U. S. Fish Comm. 11 : 409 ( 1893). 
Microsporidies Balbiani 1882. 
Order Microsporidiida Labbe in Thierreich 5: 104 (1899). 

The parasites of this order attack chiefly arthropods and fishes. They multiply 
asexually and produce serious epizootics. The spores are very minute, and the details 
of the processes by which they are formed are unknown. A polar capsule is present in 
each spore (those of Telornyxa have two polar capsules). The polar capsules are not 
visible in living material, but are revealed by treatment with alkali. 

In Kudo's monograph of this order ( 1924) , more than 150 species are treated. They 
form four families. 

Family 1. Glugeida [Glugeidae] Gurley op. cit. 409. Glugeidees Thelohan op. cit. 
174. Glugeidea Braun I.e. Family Nosematidae Labbe in Thierreich 5: 104 (1899). 
Family Plistophoridae Doflein Protozoen 205 ( 1901 ) . Spores oval, ovoid, or pyriform. 
Nosema Bombycis Nageli causes the pebrine disease of silkworms; another species of 
Nosema causes an epizootic of honeybees. Gliigea attacks several species of fishes. 
Gurleya, Thelohania, Duboscquia, Plistophora, etc. 

Family 2. Coccosporida [Coccosporidae] Kudo Handb. Protozool. 323 (1931). 
Family Cocconemidae Leger and Hesse 1922, based on a generic name which is a 
later homonym. Spores globular. Coccospora Slavinae (Leger and Hesse) Kudo, in 
the oligochaet worm Slavina. 

Family 3. Mrazekiida [Mrazekiidae] Leger and Hesse 1922. Spores elongate, 
exceedingly minute, resembling bacteria. Mrazekia, Octospora, Spironema, 

Family 4. Telomyxida [Telomyxidae] Leger and Hesse 1910. Telornyxa glugei- 
formis, in the fat body of the larva of Ephemera vulgata, producing ellipsoid spores 
with a polar capsule at each end. 

Chapter XII 

Phylum 8. CIUOPHORA (Doflein) nomen phylare novum 

Class Infusoires Lamarck Phil. Zool. 1: 127 (1809). 

Class Infusoria Lamarck Anim. sans Verteb. 1: 392 (1815). 

Class Protozoa Goldfuss in Isis 1818: 1008 (1818). 

Class Polygastrica Ehrenberg Infusionsthierchen p.* (1838). 

Hauptgruppe Protozoa, class Infusoria, and order Stomatoda Siebold in Siebold 
and Stannius Lehrb. vergl. Anat. 1 : 3, 10 ( 1848). 

Subkingdorn Archezoa Perty Kennt. kl. Lebensf. 22 (1852), not phylum Archezoa 
Haeckel (1894). 

Order Ciliata Perty op. cit. 137. 

Subphylum Infusoria Haeckel Gen. Morph. 2: Ixxviii (1866). 

Phylum Infusoria Haeckel Syst. Phylog. 1: 90 (1894). 

Subphylum Ciliophora Doflein Protozoen 227 (1901). 

Dependent organisms, mostly predatory, unicellular but mostly of complicated 
structure; swimming by means of cilia at least at some stage of life; mostly with 
nuclei of two types in each cell. Vorticella, the only genus named by Linnaeus, is to 
be considered the type. 

These organisms are the typical examples of the accepted groups Infusoria and 
Protozoa. The name Infusoria, referring to creatures which appear in infusions, is 
said to have been introduced by Ledermiiller, 1763, or Wrisberg, 1764. As a scien- 
tific name it has status from its application to a class by Lamarck (1815). The name 
Protozoa, applied to a class in its original publication by Goldfuss, is a later synonym 
of Infusoria. In treating the group as a phylum, one finds it necessary to apply a new 
name, and takes up as such the name which Doflein applied to it as a subphylum. 

The essential point in the definition of the phylum is the word cilia. Cilia are cell- 
organs of the same nature as flagella, differing in being smaller in proportion to the 
cell which bears them, more numerous, and distributed generally on the surface. In 
Loeffler's classic investigation (1889), they were found to bear solitary terminal ap- 
pendages; by subsequent terminology, they are acroneme. Doflein appears to have 
been mistaken in emphasizing the difference between flagella and cilia; there is no 
fundamental difference. A verbal distinction, nevertheless, is expedient: the applica- 
tion of the term ciHum is to be restricted to two things, (a) the swimming organelles 
of the Ciliophora, and (b) moving fibrils protruding abundantly from certain epithe- 
Hal cells of animals. Botanical usage, which treats cilium and flagellum as synonyms, 
is unsound. The structures which in botany have been called cilia are definitely 

The cells of Ciliophora reach moderately large sizes; those of the classroom 
example Paramaecium attain a length of 0.25 mm. and are perceptible to the naked 
eye. The cells of some of the Ciliophora are the most highly compHcated of all indi- 
vidual cells. In addition to the cilia, the cell organs which require discussion are the 
pellicle, neuromotor fibrils, trichocysts, structures involved in nutrition, contractile 
vacuoles, and nuclei. 

The cell has a firm ectoplasm or pellicle which gives it a definite form. The cilia 
spring from basal granules imbedded in the pellicle. In simpler examples, the cilia 

224 ] The Classification of Lower Organisms 

are essentially uniform and uniformly distributed on the surface. Other examples 
are without separate cilia upon part or all of the surface, but bear a variety of struc- 
tures which consist of coalescent cilia. Membranelles are triangular appendages con- 
sisting of brief rows of ciHa; undulating membranes represent long rows; cirri represent 
tufts. The organisms of class Tentaculifera bear cilia only in the juvenile condition. 
At maturity they bear extensible tubular structures called tentacles, by means of 
which they capture free-swimming ciUates and absorb their contents. 

The basal granules of the cilia are linked together by a system of fibrils; the cilia 
and fibrils make up the neuromotor apparatus. This term was coined by Sharp, in his 
study of Diplodinium (1914). The neuromotor fibrils form a highly elaborate net- 
work, not connected with the nucleus, as in flagellates, but to a central structure, 
apparently regulative, called the motorium. The motorium of Diplodinium is a mas- 
sive body near the anterior end; that of the tintinnids is fairly large in proportion to 
the cells (Campbell, 1926, 1927); that of Paramaecium, presumably a comparatively 
primitive organism, is a minute body lying near the dorsal side of the cytopharynx 
(Lund, 1933). 

Imbedded in the pellicle, in addition to the neuromotor fibrils, there are certain 
minute ellipsoid bodies called trichocysts. These, when the cell is irritated, discharge 
their contents in the form of elongate rods or threads. Their mechanism and effect 
are not understood. 

In most Ciliophora, each cell has a mouth and gullet; or better, since these struc- 
tures are not homologous with those of animals, a cytostome and cytopharynx. The 
cytopharynx is a more or less funnel-shaped impression in the cell. It is bounded 
laterally by ciliate pellicle; its outer opening is the cytostome; it is closed at the inner 
end by a layer of cell membrane directly over fluid cytoplasm. Prey, chiefly bacteria 
and small algae, encountered by the organism as it swims, is swept into the cyto- 
pharynx by the action of the cilia. When a certain mass of prey has accumulated, 
the cell membrane at the inner end of the cytopharynx becomes impressed and under- 
goes constriction, enclosing the prey in a food vacuole. The material in the food 
vacuole undergoes digestion; while this is taking place, movement of the cytoplasm 
carries the vacuole along a definite circuitous course through the interior of the cell. 
After some time, the vacuole arrives at a certain point on the pellicle, the anus or 
cytoproct, where it discharges its contents and disappears by bursting through the 

In freshwater species, each cell contains one or more contractile vacuoles which 
appear at definite points and disappear periodically by discharge of their contents 
to the exterior. Associated with the proper contractile vacuoles, there may be systems 
of "canals" which are in fact additional contractile vacuoles. These structures have 
been much studied; there are notable accounts by Day (1930) and Mac Lennan 
(1933). When a vacuole has disappeared by discharge, it reappears as one or more 
minute vacuoles in the same area: minute bodies of gelled protoplasm turn into sol, 
and then become lifeless liquid. The discharge of a "canal" into the proper con- 
tractile vacuole occurs by dissolution of the bounding membranes of gelled proto- 
plasm where the two are in contact, followed by contraction of the membrane of the 
canal. The proper contractile vacuole discharges by essentially the same mechan- 
ism. Its membrane meets and becomes fused with the bounding membrane of the 
cell, generally at the end of a brief channel through the pellicle; the combined mem- 
brane breaks, and the membrane of the vacuole contracts. 

Phylum Ciliophora [ 225 

Earlier biologists supposed tliat the contractile vacuole is an excretory mechanism. 
More probably, its function is purely hydrostatic, to rid the cell of the water which 
is constantly entering by osmosis. Marine Ciliophora have no contractile vacuoles. 

In many members of the family Opalinoea each cell has many similar nuclei. These 
divide, from time to time, by mitosis. Cell division takes place independently of 
nuclear division, by transverse constriction, when a certain size has been reached. 

In the generaHty of Ciliophora, each cell has one or more nuclei of each of two 
types, macronuclei, which are conspicuous, and micronuclei, discerned with difficulty. 

Cell division occurs by transverse constriction and is necessarily associated with 
nuclear division. The macronucleus becomes elongate and divides by constriction 
without any formation of chromosomes; in other words, amitotically. The micro- 
nucleus also becomes elongate and divides by constriction. Early observers supposed 
this process also to be amitotic. Actually, there appear within the intact nuclear 
membrane a spindle and a definite number of chromosomes. Reichenow (editing 
Doflein, 1927) compiled the following diploid counts: 

Stentor coeruleus 28 

Didinium nasutum 16 

Chilodon uncinatus 4 

C arche Slum poly pinum 16 

Turner (1930) found 8 in Euplotes Patella. Thus the chromosome numbers of 
Cihophora appear usually to be small powers of 2. 

The chromosomes duly undergo division, the daughter chromosomes going to dif- 
ferent ends of the nuclear cavity. The nucleus becomes greatly elongate and its mem- 
brane presses in from the sides and cuts it in two. Turner observed in the axis of the 
spindle of Euplotes a rather small endosome which becomes elongate and undergoes 
constriction while the chromosomes are forming. 

Opalina has a sexual process in which the multinucleate cells divide into many 
uninucleate gametes. These are sexually differentiated, larger and smaller; they 
duly unite in pairs and the zygotes grow and become ordinary multinucleate 

In the generality of Ciliophora, early observers discovered a sexual process in 
which the cells, apparently undifferentiated, join in pairs but maintain their individ- 
uality. The uniting cells become attached to each other in definite positions: in 
Paramaecium, by their ventral or mouth-bearing surfaces; in Euplotes, by the left 
halves of their broad ventral surfaces; in the ophryoscolecids and various other groups, 
by their anterior ends. They remain attached, while continuing to swim, for several 
hours, during which an exchange of nuclei takes place, and then resume their separate 
life. Calkins (1926) was disposed, contrary to historical usage, to confine application 
of the term conjugation to this exceptional form of syngamy. 

The nuclear details of conjugation were described by Maupas (1889) and Richard 
Hertwig (1889), whose observations have repeatedly been confirmed. When a pair 
have joined, their macronuclei divide several times; the ultimate fragments are 
digested and disappear. The micronuclei also divide, concurrently in both conju- 
gants, a fixed number of times, in Paramaecium three, in Euplotes four. These divi- 
sions include a meiotic process. Most of the haploid nuclei produced are digested; 
as a general rule, only one survives to undergo the final division, which is mitotic, 
producing in each conjugant two genetically identical haploid nuclei. By this time 
a cytoplasmic connection has been established between the conjugants. In Paramae- 
cium, the spindles of the mitotic final nuclear divisions extend through this connection, 

226 ] The Classification of Lower Organisms 

so that when mitosis is complete each protoplast contains two haploid nuclei of dif- 
ferent origin. In other ciliates the same result is attained, apparently, by the migration 
of one nucleus of each pair. Karyogamy takes place in each conjugant. The cyto- 
plasmic connection is broken and the conjugants separate from each other. During 
several subsequent hours, the zygote nucleus undergoes a characteristic number of 
divisions, three in Paramaecium. Among the nuclei produced, one usually enlarges 
and becomes a macronucleus; others, of the number characteristic of the form, survive 
as micronuclei; the remainder are digested. 

In Vorticella and its allies, syngamy consists of the complete fusion of a smaller 
swimming individual with a larger one attached by a stalk. The nuclear processes 
are believed to be essentially as in other ciliates. The reproduction of the Tentaculi- 
fera has not been much studied, but here also the nuclear changes are as in the 
generahty of ciliates (Noble, 1932). 

The possibility of conjugation is limited by the occurrence of mating types. Certain 
early observations had suggested the existence of these; the definite discovery was by 
Sonneborn, in Paramaecium Aurelia (1937). Results of further study are available 
in a symposium edited by Jennings (1940) and in a review by Kimball (1943). To 
current knowledge, then: 

Paramaecium caudatum includes four mating types divided into two groups; types 
I and II conjugate with each other, and types III and IV with each other, but the 
two groups are mutually sterile. 

Paramaecium Aurelia includes eight mutually sterile groups, each of two mutually 
fertile mating types. 

Paraviaecium Bursaria includes three mutually sterile groups. The first group is 
of four types, each self-sterile but able to conjugate with any other; the second group 
is of eight such types, and the third again of four. 

Paramaecium. multimicronucleatum is without mating types; any race can conju- 
gate with any other 

Euplotes Patella includes six mating types all in one group; each can conjugate 
with any other. 

The heredity of mating types is not understood. It is not a matter of simple Men- 
dclian heredity. In Paramaecium Bursaria group I, the progeny of a cell of a given 
mating type may include after conjugation either two or all four of the mating types. 
The mating type of a line becomes fixed in connection with the first or second cell 
division after conjugation, at the time that macronuclei are being differentiated; it 
is accordingly believed that something in the macronuclei fixes the mating types. 

So far as mating types are present, pure lines of ciliates cannot conjugate. Early 
attempts to maintain pure cultures failed by death after intervals of some months. 
These observations led to speculations that the vitality of protoplasm is limited, and 
that sexual reproduction restores it. Woodruff, however, proved it possible to maintain 
Parflmagau?n ^urc/m indefinitely without conjugation: he reported (1926) a culture 
so maintained for sixteen years, an estimated eleven thousand generations. 

The cultures are not thus persistent without nuclear change. At intervals, the macro- 
nuclei break up and dissolve, and are replaced by new ones formed by division of the 
micronuclei. Woodruff and Erdmann (1914) applied to this process of replacement 
of nuclei the term cndomixis. It is not possible that this process is the genetic equiva- 
lent of karyogamy. It is, presumably, the physiological equivalent of conjugation in 
its feature of providing new macronuclei. 

Phylum Ciliophora [ 227 

Diller (1936) observed in P. Aurelia a different manner of replacement of nuclei, 
by autogamy. In this process, the nuclei of a solitary cell go through the preliminaries 
of conjugation; two haploid nuclei, sister products of one act of mitosis, unite to form 
a zygote nucleus; and this divides in the usual manner to produce micronuclei and 
macronuclei. Wichtermann (1939, 1940) observed that two cells, joined as in conju- 
gation, may simultaneously undergo autogamy instead of exchanging nuclei. 

In the normal conjugation of ciliates, the gamete nuclei produced in each cell, 
being sister products of mitosis, are genetically identical; and the zygote nuclei pro- 
duced after interchange are also genetically identical with each other. Autogamy is 
believed to produce diploid nuclei which are completely homozygous. Thus the sexual 
processes of the ciliates tend strongly to limit the variability of the progeny. This is a 
peculiar and surprising feature of the group. 

The ciliates have attracted experimental study, beyond what has already been 
implied, of various functions, including nutrition, inheritance of acquired characters, 
and regeneration after injury. 

Hall and his associates (1940-1945) have shown that Colpidiinn campylum and 
Tetraphymena Geleii (the latter is in their earlier papers called Glaucoma piriformis) 
require thiamin and probably riboflavin. Nutritional requirements, rather than such 
an entity as vitality, are presumably responsible for the limited life of early attempted 
pure cultures. As to minerals, the same scholars demonstrated the necessity of Ca and 
Fe: others have demonstrated the necessity of K, Mg, and P. 

It has been observed of certain cultures in which the rate of division has been in- 
creased by exposure to high temperature that they would continue to divide abnor- 
mally rapidly when returned to normal temperatures. The peculiarity disappeared in 
individuals which conjugated. By refrigeration or by application of chemicals, there 
have been produced "monsters," individuals of abnormal structure, which have repro- 
duced themselves through many generations, and have proved capable eventually of 
giving rise to normal individuals. Jollos (1913) designated as Dauermodifikationen, 
that is, enduring changes, modifications of the type described. They are actually 
acquired characters which can be inherited within limits. It is evident that they are 
determined by macronuclei or by cytoplasm, and that they are not in conflict with 
the principle that the truly enduring heredity of nucleate organisms Hes in nuclei 
which divide mitotically. 

Balamuth (1940) reviewed the literature of experimental mutilation of Protozoa 
and gave a bibliography of 173 titles. Most of the experiments have been performed 
on ciliates. The conclusions from them include these, that regeneration of parts arti- 
ficially cut away takes place with different degrees of facility in different groups, 
and that it is effected, if at all, by the same mechanism by which the parts are pro- 
duced after division or excystment. The less elaborate ciliates, as Opalina and 
Paramaecimn, are usually killed by mutilation, since this allows the fluid inner 
cytoplasm to escape. In Stentor, injury to the crown of membranelles results in the 
appearance of a new crown of membranelles on the side of the body, followed by its 
migration to the injured area. In Stylonychia and Euplotes, destruction of one cirrus 
is followed by the appearance, in a certain area of the surface, of the primordia of a 
complete set of cirri; the original cirri are absorbed, and the new ones migrate along 
the surface to their proper stations. The regulation of regeneration is explained, as 
are various other phenomena, in a review by Weisz (1954). 

Micronuclei are necessary for unlimited hfe and for sexual reproduction, but not 
for regeneration and a long period of Hfe. Schwartz kept a culture of Stentor alive 

228 ] The Classification of Lower Organisms 

without micronuclei for more than a year. Macronuclear material is necessary for 
regeneration, but any fragment of a macronucleus is sufficient. This is a very signifi- 
cant observation. It means that all the factors controlling the vegetative structure and 
behavior of a cell can be spread out and intermingled in all parts of a body of con- 
siderable size; it furnishes an analogy to the state of affairs which may be supposed 
to exist in bacteria. 

The Ciliophora are treated as two classes, Infusoria and Tentaculifera. Hartog 
(1909) estimated the number of known species of the former as about five hundred. 
This number would have included practically all of the fresh-water species known up 
to the present. Entozoic and marine species were known, but hundreds of species of 
these ecological groups have subsequently been discovered. Including some two 
hundred species of Tentaculifera, the phylum Ciliophora appears to be of about 
twelve hundred known species. 

Class 1. INFUSORIA Lamarck 

Class Ciliata Haeckel Gen. Morph. 2: Ixxviii (1866). 
Class Ciliatea Hall Protozoology 333 (1953). 
Further synonymy essentially as of the name of the phylum. 

Ciliophora lacking tentacles, bearing cilia or modified cilia in the mature condition. 

Stein (1867) provided four orders of Infusoria. These orders are surely natural. 

Subsequent authors have proposed many modifications of Stein's system, and many 

of these are surely sound; but among groups proposed as additional orders, only the 

opahnids are positively entitled to this status. 

1. Nuclei all alike, commonly numerous. Order 1. Opalinalea. 

1. Nuclei diflferentiated into macronuclei and 

2. Without a spiral band of membranelles 

or cilia about the cytostome Order 2. Holotricha. 

2. With a spiral band of membranelles or 
cilia about the cytostome. 
3. The spiral sinistrorse. 

4. Not of the character of the fol- 
lowing order Order 3. Heterotricha. 

4. Flattened, cirri and most cilia 

confined to the ventral surface Order 4. Hypotricha. 

3. The spiral dextrorse Order 5. Stomatoda. 

Order 1. Opalinalea nom. nov. 

Suborder Opalininea Poche in Arch. Prot. 30: 250 (1913). 

Protociliata Metcalf in Anat. Record 14: 89 (1918) and Jour. Washington Acad. 

Sci. 8: 431 (1918). 
Subclass Protociliata Kudo Handb. Protozool. 335 (1931). 
Order Opalinida Hall Protozoology 113 (1953), preoccupied by family Opalini- 
dae Claus. 
Nuclei not differentiated into two types; cilia abundant, undifferentiated; sexual 
reproduction by the complete union of differentiated minute uninucleate gametes. 
Commensal in the gut of amphibia and fishes. 

The group has been treated monographically by Metcalf (1923). A single family 
is usually recognized. 

Phylum Ciliophora [ 229 

Family Opalinoea Pritchard 1842. Family Opalinaea Siebold in Siebold and 
Stannius Lehrb. vergl. Anat 1: 10 (1848). Family Opalinina Stein Org. Inf. 2: 169 
(1867). Family Opalinidae Glaus 1874. Family Protoopalinidae Metcalf 1940. 
There are about 150 known species of four approximately equally numerous genera: 
Protoopalina Metcalf, cylindrical, with one or two nuclei which are always found in 
a stage of mitosis; Zelleriella Metcalf, similar, the cells flattened; Cepedia Metcalf, 
cylindrical, with many nuclei; Opalina Purkinje and Valentin, flattened and 

Order 2. Holotricha Stein Org. Inf. 2 : 169 ( 1867) . 

Orders Gymnostomata and Trichostomata, and suborder Aspirotricha Biitschli 

in Bronn Kl. u. Ord. Thierreichs 1: 1674 (1889). 
Suborder Hymenostomata Hickson 1903. 
Orders Gymnostomataceae and Aspirotrichaceae Hartog in Cambridge Nat. 

Hist. 1: 137 (1909). 
Order Holotricha with suborders Anoplophryinea, Gymnostomata, and Hymeno- 
stomata Poche in Arch. Prot. 30: 250-255 (1913). 
Order Holotrichida Calkins Biol. Prot. 376 (1926). 
Infusoria with differentiated macronuclei and micronuclei, with simple cilia dis- 
tributed generally over the surface of the body, not having membranelles in a spiral 
band about the cytostome. 

This group is the mass of the more primitive typical Infusoria, of numerous fami- 
lies, not all of which are to be listed here. Arrangements of the families in other 
groups than the three here maintained have been proposed and are presumably more 
nearly natural. 

a. Cytostome anterior. Suborder Gymnostomata (Biitschli) Poche. Suborder stom,ina Hall. 

Family Enchelia Ehrenberg Infusionsthierchen 298 (1838). Family Enchelina 
Stein Org. Inf. 2: 169 (1867). Family Enchelyidae Kent. Families Holophryidae and 
Cyclodinidae Schouteden. Family Didiniidae Poche. Comparatively unspecialized 
forms, radially symmetrical or nearly so. Enchelis O. F. Miiller; Holophrya, Chaenia, 
Prorodon; Ichthiophthirius, becoming parasitic in the skins of fishes; Lacryviaria, 
the cytostome at the end of an extensible proboscis; Didinium, barrel-shaped, with 
the cilia confined to two belts, having an extensible proboscis by means of which it 
seizes other Infusoria and through which it swallows them. 

Family Colepina Ehrenberg op. cit. 316 includes the single genus Coleps. The cells 
look like hand grenades of World War I: they are approximately barrel-shaped (the 
axis more or less curved), the pellicle forming hardened quadrangular plates between 
which the cilia project. The anterior cytostome can be opened widely to ingest other 

b. Cytostome lateral. Suborder Aspirotricha Biitschli. 

Family Parameciina Perty (1852). Family Paramoecidae Grobben. Paramaecium 
[Hill] O. F. Miiller Verm. Terr. Fluv. 1: 54 (1773). The name is variously spelled; 
the spelling here used is Miiller's in what is believed to be the first publication under 
binomial nomenclature. 

Family Colpodaea Ehrenberg Infusionsthierchen 345 (1838). Family Colpodidae 
Glaus 1879. Family Ophryoglenidae Kent 1882. Small forms, oval, bean-shaped, 
or flattened. Ophryoglena, Glaucoma, Colpoda, Tetrahymena, and many others. 

230 ] The Classification of Lower Organisms 

Family Cyclidina Ehrenberg op. cit. 244. Family Pleuronemidae Kent. Family 
Pleuronemina Biitschli (1889). Similar, with a conspicuous undulating membrane 
along one side. Cyclidium and many other genera. 

Family Urocentrina Claparede and Lachmann Etudes Inf. 1 : 134 ( 1858) . Family 
Urocentridae Schouteden. Urocentrum, the single genus, top-shaped, with cilia con- 
fined to two belts and a tail-like tuft, constantly whirling in the water. 

Family Trachelina Ehrenberg op. cit. 319. Family Tracheliidae Kent. Having an 
anterior proboscis, the mouth at the base of this. Trachelius, Dileptus, Lionotus, 
Loxodes, etc. 

Family Chlamydodontida [Chlamydodontidae] Glaus 1874. Family Chlamydo- 
donta Stein, the mere plural of a generic name. Family Chilodontida Biitschli. Fam- 
ily Nassulidae Schouteden. Flattened. The cytopharynx surrounded by longitudinal 
rods, apparently of hardened protein, collectively forming a conical basket, enclosed 
except when the cytostome is open for ingestion. Chilodon, Chlamydodon, Nassula. 
c. Cytostome lacking; parasitic, mostly in invertebrates. Suborder Anoplophry- 
INEA Poche; suborder Astomina Hall. 

Family Anoplophryida [Anoplophryidae] and seven other families, all named by 
Cepede, 1910. 

Order 3. Heterotricha Stein Org. Inf. 2: 169 (1867). 

Suborder Spirotricka, sections Heterotricha and Oligotricha Biitschli in Bronn 

Kl. u. Ord. Thierreichs 1 : 1674 ( 1889). 
Section Chonotricha Wallengren in Acta Univ. Lund 31, part 2, no. 7 : 48 ( 1895) . 
Order Oligotricha Doflein Protozoen 240 (1901). 
Orders Heterotrichaceae and Oligotrichaceae Hartog in Cambridge Nat. Hist. 

1: 137 (1909). 
Orders Heterotrichida and Oligotrichida Calkins Biol. Prot. 386, 388 (1926). 
Suborder Entodiniomorpha Reichenow in Doflein Lehrb. Prot. ed. 5, 3: 1195 
(1929); Order Chonotricha Reichenow op. cit. 1211; suborder Ctenostomata 
Kahl ex Reichenow op. cit. 1024. 
Orders Spirotrichida and Chonotrichida Hall Protozoology 380, 411 (1953). 
Infusoria having a sinistrorsc spiral band of cilia about the cytostome, these cilia 
united (except in family Spirochonina) in triangular-attenuate membranellcs; not 
having the body flattened and the cilia or cirri confined to the ventral surface. 

The peristomal apparatus of this order is an evidently derived character, so pecu- 
liar as to appear to have evolved only once: in short, the order appears natural. There 
are numerous subordinate groups. Several of these, of many species or of exceptional 
character, have been segregated as additional orders; it is by an arbitrary decision 
that they are here treated as suborders. 

a. Comparatively unspecialized examples. Suborder Spirotricha Biitschli. 
Suborders Hctcrolrichina and Oligolrichina Flail. 
Family Plagiotomina Biitschli op. cit. 1719 (1889). Family Plagiotomidae Poche 
(1913). Peristomal area narrow and elongate, extending from the anterior end to a 
cyto.^tome located near the middle of one side. Blepharisyna. Spirostomum. 

Family Bursarina Stein Org. Inf. 2: 169, 295 (1867). Family Bursariidae Kent. 
Cytostome seated in a deep pit in one side of the body. Bursaria. Balantidium, para- 
sitic in the gut of Amphibia and mammals; B. coli, a serious pathogen in man. 

Family Stentorina Stein op. cit. 169, 217. Family Stentoridae Claus. Peristomal 
area anterior, more or less transverse. Stcntor, sessile and obconic, familiar. Follicu- 

Phylum alio phor a [231 

Una, the posterior end seated in a chitinous lorica, the peristomal area broadly ex- 
panded as two wings. 

Family Halterina Claparede and Lachmann Etudes Inf. 1: 367 (1858). Family 
Halteriidae Claus. Halteria, subglobular, with a single whorl of long cilia; familiar in 
infusions, recognizable by the motion of the cells, alternately revolving slowly and 
snapping violently from place to place. 

b. Loricate, free-swimming. Suborder Tintinnoinea Kofoid and Campbell. 
Suborder Tintinnina Hall. 

Family Tintinnodea Claparede and Lachmann Etudes Inf. 1: (1858). Family 
Tintinnidae Claus. Peristomal membranelles elongate and ciliate, the cylindrical or 
conical body attached in and retractile into the lorica; characteristically with two 
macronuclei and two micronuclei. Mostly marine. Kofoid and Campbell, who mono- 
graphed the group ( 1929) , found it possible to distinguish the natural and subordinate 
groups entirely by the structure of the lorica. They divided the former single family 
into twelve and recognized more than three hundred species. 

c. Laterally flattened, with a tough membrane and few cilia and membranelles. 
Suborder Ctenostomata Kahl. Suborder Ctenostomina Hall. 

Family Ctenostomida [Ctenostomidae] Lauterborn in Zeit. wiss. Zool. 90: 665 
(1908). Kahl (1932) monographed the group and found twenty-five species, which 
he arranged in six genera and three families. 

d. Cylindrical, entozoic, with no ciliation except the membranelles. Suborder 
Entodiniomorpha Reichenow. Suborder Entodiniornorphina Hall. 

Family Ophryoscolecina Stein Org. Inf. 2: 168 (1867). Family Ophryoscolecidae 
Claus. Becker (1932) reviewed previous studies of this group, examples of which 
were first mentioned by Gruby and Delafond, 1843. He noted 71 species, of the 
genera Entodinium, Diplodinium, Ophryoscolex, Epidinium, etc. (the genera were 
first named by Stein) in the domestic ox; and 52 {Didesmis, Paraisotricha, Spirodin- 
ium, Cycloposthium, etc.) in the horse. Dogiel (1927) monographed the family, but 
it is certain that large numbers of species remain to be discovered in wild animals, 
oxen and others. 

The barrel-shaped cells are about 0.1-0.25 mm. long. The cytostome is anterior, 
surrounded by the usual spiral band of membranelles; this may be broken up into 
several partial files, and there may be belts or clusters of membranelles on other parts 
of the body. The posterior end is drawn out into processes, one, few, or many, ob- 
scure or prominent, horn-like or fringe-like. Internally, beside contractile vacuoles 
and a neuromotor apparatus including a large motorium, there are characteristic 
skeletal plates. These consist of minute cylindrical bodies imbedded in an amorphous 
matrix, the whole staining with iodine and consisting supposedly of some polysac- 
charide carbohydrate. 

Animals are infected by eating food contaminated with the saliva of others. The 
ciliates may be present in the rumen in numbers from one thousand to three million 
per cc. It has been supposed that they are symbiotic, benefitting their hosts by carry- 
ing on useful syntheses, or perhaps merely by controlling numbers of bacteria in the 
rumen. There is no good evidence for these ideas: the probability is, that they are 
harmless commensals. 

e. Cylindrical or obconic, sessile, cilia of the peristomal band separate, body 
otherwise naked. Suborder Chonotricha (Wallengren) subordo novus. 

Family Spirochonina Stein Org. Inf. 2: 168 (1867). Family Spirochonidae Grob- 


The Classification of Lower Organisms 

Fig. 44. — Infusoria, order Hypotricha: a, Aspidisca x 800. b, Stylonychia 
X 400. C, Euplotes x 400. d-n, Euplotes Patella after Turner (1930); d-h, stages 
of mitosis x 2,000, i, conjugating cells x 400; j, k, polar and equatorial views of the 
heterotypic division in a conjugant x 2,000; 1, early anaphase of the homeotypic 
division x 2,000; m, first division of the zygote nucleus x 2,000; n, a cell after con- 
jugation X 400, the macronucleus breaking up, the zygote nucleus divided into four, 
of which one is to become a macronucleus, one a micronucleus, and two are to 
undergo dissolution. 

Phylum Ciliophora [ 233 

ben. Spirochona and a few other genera, attached to aquatic animals, fresh-water or 
marine, best known from the crustacean Gammarus. 

Order 4. Hypotricha Stein Org. Inf. 2 : 168 ( 1867) . 

Section Hypotricha Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 1674 (1889). 
Order Hypotrichaceae Hartog in Cambridge Nat. Hist. 1: 137 (1909). 
Order Hypotrichida Calkins Biol. Prot. 389 (1926). 
Suborder Hypotricha Kudo Man. Protozool. ed. 3: 668 (1946). 
Suborder Hypotrichina Hall Protozoology 381 ( 1953 ) . 
Flattened Infusoria bearing a band of membranelles crossing the upper surface 
near the anterior end from right to left and continued rearward on the lower surface 
beside the cytostome, along which lie also undulating membranes; mostly bearing 
cirri, which are confined to the lower surface, as are most free cilia, if these are 

This group is evidently natural, and evidently a specialized offshoot from the pre- 
ceding order. It might reasonably be treated as a subordinate group of the preceding 
order; Biitschli, Kudo, and Hall have done so. There are comparatively few species. 
Several are familiar in infusions and have been much studied. 

Family 1. Peritromina Stein Org. Inf. 2: 168 (1867). Family Peritromidae Kent. 
Cilia abundant on the lower surface, cirri none. Peritromus. 

Family 2. UrostyUda [Urostylidae] Calkins Biol. Prot. 390 (1926). As above, but 
with frontal and sometimes also anal cirri. Numerous genera, Urostyla, Uroleptus, 
Epiclintes, Stilotricha; Kerona O. F. Miiller, an ectoparasite on the animal Hydra. 
Family 3. Oxytrichina Ehrenberg Infusionsthierchen 362 (1838). Family Oxytri- 
chidae Kent. Family Pleurotrichidae Biitschli. Cirri present; cilia in one or two mar- 
ginal rows, few or absent on the ventral surface. Oxtricha, Stylonychia, Pleurotricha, 
Euplotes, etc. 

Order 5. Stomatoda Siebold in Siebold and Stannius Lehrb. vergl. Anat. 1: 10 

Order Ciliata Perty Kennt. kl. Lebensf. 137 (1852). 

Order Peritricha Stein Org. Inf. 2: 168 (1867), 

Section Peritricha Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 1674 (1889). 

Order Peritrichaceae Hartog in Cambridge Nat. Hist. 1: 138 (1909). 

Order Peritrichida Calkins Biol. Prot. 395 (1926). 
Infusoria having a dextrorse spiral band of membranelles about the cytostome, 
which can in most examples be concealed and protected by contraction of the body; 
free-swimming only in the immature condition, at maturity attached and without 
separate cilia; syngamy occurring by the complete union of a smaller swimming indi- 
vidual with a larger attached one. Vorticella is the apparent type of the old ordinal 
names Stomatoda and Ciliata, which are accordingly held to belong to this order. 

Family Vorticellina Ehrenberg Infusionsthierchen 259 (1838). Family Vaginifera 
Perty (1852). Family Vorticellidae Fromentel 1874. Vorticella L., a familiar mic- 
roscopic organism in material from ponds and ditches, consists of solitary bell-shaped 
cells on contractile stalks. Carchesium and Zoothamnium are similar organisms in 
colonies. Ophrydium, Epistylis, etc., consist of similar colonies of non-contractile 
cells. Cothurnia and Vaginicola are solitary stalkless cells having conical loricae into 
which they can withdraw themselves. 


The Classification of Lower Organisms 

Fig. 45 — Tokophyra Lemnarum after A. Noble (1932) : a, representative individ- 
ual; b, budding; c, swimming bud; d, conjugation; e, feeding on a cell of Euplotes; 
t, cyst; g, tentacles, feeding, expanded, and contracted, g x 2,000, all others x 400. 

Phylum Ciliophora [ 235 

Family Urceolarina Perty (1852). Family Trichodinidae Glaus. Family Urceol- 
aridae Kudo. Urceolaria, Trichodina, etc., disk- or barrel-shaped cells attached on 
or in aquatic animals by means of a whorl of hard hooks. 

Class 2. TENTACULIFERA (Huxley) Kent 

Order lufusoires suceurs and group Acinetina Claparede and Lachmann Etudes 
Inf. 1: 377,381 (1858). 

Class Acinetae Haeckel Gen. Morph. 2: Ixxix (1866), the mere plural of a generic 

Tentaculifera Huxley Man. Anat. Invert. 100 (1877). 

Glass Tentaculifera with orders Suctoria and Acinetaria Kent Man. Inf. 1 : 36 

Class Acinetaria and order Suctoria Lankester in Enc. Brit. ed. 9, 19: 865 ( 1885). 

Subclass Suctoria Butschli in Bronn Kl. u. Ord. Thierreichs 1: 1842 (1889). 

Class Acinetoidea Poche in Arch. Prot. 30: 263 (1913). 

Class Sudor ea Hall Protozoology 413 (1953). 

Organisms swimming by means of cilia while immature, at maturity lacking cilia 
and usually attached, provided with tentacles by which they capture and paralyze 
their prey and absorb food. Acineta is the type genus. 

These organisms are rather unfamiliar. They occur both in fresh water and in salt, 
and prey chiefly upon Infusoria. There are differentiated macronuclei and micro- 
nuclei; in branching or colonial individuals, a single macronucleus may extend to all 
parts. Asexual reproduction is by budding, often endogenous. Conjugation occurs 
either between attached individuals or between an attached individual and a swim- 
ming bud. The fact that one individual may bend past another to conjugate with a 
third indicates the presence of mating types. Conjugating individuals exhibit pregamic 
and postgamic nuclear divisions quite as among Infusoria (Noble, 1932). The group 
is undoubtedly derived from Infusoria; whether from something of the nature of 
Didinium, Vorticella, or Spirochona remains uncertain. 

Collin (1912) accounted for about 170 species and recognized eight families. One 
of these families has subsequently been transferred to order Holotricha. The re- 
mainder may be construed as a single order: 

Order Suctoria Kent (1880). Lankester chose this as between two ordinal names 
which Kent published at the same time. 

a. Individuals subglobular, usually stalked, their tentacles essentially uniform. 
Family 1. Podophryina Butschli in Bronn Kl. u. Ord. Thierreichs 1 : 1926 (1889). 

Family Podophryidae, Rousseau and Schouteden 1907. Buds produced exogenously. 
Podophrya, Sphaerophrya, naked; Urnula, loricate. 

Family 2. Acinetida [Acinetidae] Glaus 1874. Acinetina Claparede and Lach- 
mann (1858). Family Acinetina Biitschli (1889). Bodies with a thin pellicle, with or 
without loricae; budding endogenous. Acineta, Tokophrya, etc. 

Family 3. Discophryida [Discophryidae] Collin in Arch. Zool. Exp. Gen. 51: 364 
(1912). Body with a firm pellicle, budding endogenous. Discophrya, etc. 

b. Individuals branching or colonial. 

Family 4. Dendrosomida [Dendrosomidae] Kent Man. Inf. 2: 215 (1882). Family 
Dendrosomina BiitschU (1889). Family Dendrosomatidae Poche (1913). Dendro- 
soma, etc. 

236 ] The Classification of Lower Organisms 

Family 5. Ophryodendrida [Ophryodendridae] Kent I.e. Family Ophryodendrina 
Biitschli (1889). Ophryodendron, etc. 

Family 6. Dendrocometida [Dendrocometidae] Kent I.e. Family Dendrocometina 
Biitschli (1889). Dendrocometes, Stylocometes. 

c. With differentiated tentacles for piercing and sucking. 

Family 7. Ephelotida [Ephelotidae] Kent. I.e. Family Ephelotina Sand 1899. 
Marine, individuals subglobular, stalked. Ephelota, naked; Podocyathus, loricate. 

With this peculiar and highly evolved group, the here-proposed classification of 
organisms which lack the distinctive characters both of plants and of animals is 

List of N omenclatural Novelties [ 237 



P'amily Kurthiacea fam. nov 21 

Family Pasteurellacea nom. nov 22 

Family Chromatiacea nomen familiare novum 31 

Family Rhodobacillacea nom. nov 31 

Family Chlorobiacea nom. nov 31 

Order Sphaerotilalea nom nov 33 

Lagenocystis, nom. nov., and L. radicicola, comb, nov 82 

Family Dinamoebidina nom nov 101 

Phylum Opisthokonta phylum novum 110 

Chilomastix hominis comb, nov 165 

Pentatrichomonas obliqua comb, nov 167 

Goussia Schubergi comb, nov 207 

Family Myxoceratida and Myxoceros, nomina nova; M. sphaerulosa and 

M. Blennius, combinationes novae 221 

Phylum Ciliophora nomen phylare novum 223 

Order Opalinalea nom. nov 228 

Suborder Chonotricha subordo novus 231 


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Absidia, 123, 124 
Acantharia, 189, 190, 195, 196, 197 
Acanthochiasma, 197 
Acanthocystida, 191, 193 
Acanthocystidae, 193 
Acanthocystis, 193 
Acanthometra, 195, 197 
Acanthometren, 197 
Acanthometrida, 197 
Acanthometron, 197 
Acanthonida, 197 
Acanthophracta, 195, 197 
Acanthorhynchus, 218 
Acanthospora, 218 
Acanthosporida, 218 
Acanthosporidae, 218 
Acaulopage, 124 
Acephalina, 215 
Acervulina, 187 
Acervulinida, 187 
Acetobacter aceti, 24 
Acetobacteriacea, 20, 24 
Acetobacteriaceae, 24 
Achlya, 70, 79 
Achlya caroliniana, 78 
Achlyogeton, 118 
Achlyogetonacea, 115, 117 
Achlyogetonaceae, 117 
Achnanthea, 76 
Achnantheae, 76 
Achnanthes, 76 
Achnanthaceac, 76 
Achromatiacea, 33 
Achromatiaceae, 33 
Achromatium, 33 
Achromatium oxaliferum, 32, 33 
Achromobacter, 22 
Achromobacteriacea, 19, 21 
Achromobacteriaceae, 2 1 
Acineta, 235 
Acinetae, 235 
Acinetaria, 235 
Acinetida, 235 
Acinetidae, 235 
Acinetina, 235 
Acinetoidea, 235 
Acnidosporidea, 207 
Acrasidae, 203 
Acrasina, 203 
Acrasis, 203 
Acrita, 37 
Acrochaetiacea, 47 
Acrochaetiaceae, 47 
Acrochaetium, 47 
Actinelius, 197 
Actinollida, 197 

Actiniscea, 61, 62 

Actinisceae, 62 

Actinocephalida, 217, 218 

Actinocephalidae, 218 

Actinocephalus, 218 

Actinolophus, 193 

Actinomma, 195 

Actinomma Asteracanthion, 196 

Actinomonadida, J 91 

Actinomonadidae, 191 

Actinomonas, 190, 193 

Actinomyces Bovis, 25 

Actinomycetaceae, 25 

Actinomycetalea, 18, 24 

Actinomycetales, 24 

Actinomyxida, 219, 221 

Actinophryida, 191, 193 

Actinophryidae, 193 

Actinophrys, 193 

Actinophrys Sol, 193 

Actinopoda, 189 

Actinopodea, 189 

Actinosphaerium, 193 

Actinosphaerium Eichhornii, 192, 193 

Actipylea, 195, 197 

Actipyleen, 195 

Actipylida, 197 

Actipylina, 197 ' 

Acystosporidia, 190 

Acyttaria, 179 

Adelea, 211 

Adeleida, 211,212 

Adeleidae, 211 

Adeleidea, 211 

Adeleina, 21 1 

Adeleoidae, 21 1 

Adelina, 211 

Adinida,_98, 99 

Adiniferidea, 96, 98 

Aecidium, 147 

Aerobacter aerogenes, 22 

Agaricacea, 151, 153 

Agaricaceae, 150, 151 

Agaricales, 150 

Agaricini, 151 

Agaricus campestris, 119, 145, 152, 153 

Agarics, 152 

Aggregata, 209 

Ae;gregata Eberthi, 210 

Aggregatida, 210, 212 

Aggregatidae, 210 

Aglaozonia, 88 

Agrobacterium, 23 

Agrobactcrium tumefaciens, 23 

Agrostis, 148 

Agyriales, 137 


The Classification of Lower Organisms 

Agyrium, 137 

Ahnfeldtia, 49 

Akinetocystida, 216 

Akinetocystidae, 216 

Akinetocystis, 216 

Akinetosporeae, 86 

Albuginacea, 80, 81 

Albuginaccae, 81 

Albugo, 80, 81 

Albugo Bliti, 80 

Albugo Tragopogonis, 80 

Alcaligenes fecalis, 22 

Aleuria rutilans, 136 

Algae, 9, 10, 69, 113, 118, 120, 177, 224 

Algae, blue-green, 2, 3, 12, 13, 14, 17, 30, 

37,41, 117, 118 
Algae, brown, 39, 69, 53, 179, 203 
Algae, green, 38, 41, 53, 69, 82, 117, 118, 

128, 203 
Algae, red, 37, 39, 41, 82, 128, 140 
Algae Zoosporeae, 86 
Algen, 29, 120 
Allantocystida, 216 
Allantocystidae, 216 
Allan tocystis, 216 
Allogromia, 183 
Allogromiida, 183 
Allogromiidae, 183 
Allomorphina, 187 
Allomyces, 111, 112, 113, 118 
Allomyces anomalus, 116 
Allomyces Arbuscula, 112, 115 
Allomyces cystogenes, 112 
Allomyces javanicus, 112, 114 
Almond, 141 
Alveolina, 185 
Alveolinea, 185 
Alveolinella, 185 
Alveolinellidae, 185 
Alveolinida, 185 
Alveolinina, 185 
Alternaria, 142 
Alwisia, 175 
Amanita, 152 
Amanita muscaria, 152 
Amaurochaetacea, 174, 175 
Amaurochaetaceae, 175 
Amaurochaete, 175 
Amaurochacteae, 171 
Amaurochaetidae, 175 
Amaurosporales, 171 
Amiba diffluens, 37, 157, 201 
Amiba divergcns, 202 
Ammodiscida, 185 
Ammodiscidae, 185 
Ammodiscus, 185 
Ammodochidae, 62 
Amoeba, 71, 118, 124, 157, 189, 201 
Amoeba Proteus, 202 
Amorbaca, 201 
Amocbida, 10,201 
Amocbidac, 10, 201 

Amoebina, 201 

Amoebodiniaceae, 101 

Amoebogeniae, 219 

Amoebosporidia, 215 

Amoebosporidies, 215 

Amoebosporidiidae, 215 

Amorphoctista, 37 

Amorphozoa, 37 

Amphibia, 220 

Amphiacantha, 219 

Amphiamblys, 219 

Amphidinium, 101 

Amphilonche, 197 

Amphilothida, 103 

Amphimonadaceae, 61, 158 

Amphimonadidae, 61 

Amphisolenia, 103 

Amphisolenia laticincta, 104 

Amphistegina, 187 

Amphistomina, 191 

Anabaena, 35 

Anabaena circinnalis, 13 

Anabaena inaequalis, 32 

Ancylistales, 81 

Ancylistes, 125 

Ancylistinaeae, 81 

Anemeae, 171 

Angiococcus, 28 

Angiogastres, 152 

Angiospermeae, 82, 91 

Animacule, 18 

Animal kingdom, Animalia. Animals, 1, 2, 
4, 6, 10,' 68, 95, 111, 113, 159, 163, 
167, 206, 214, 220, 223, 231, 233, 235 

Anisochytridiales, 69 

Anisochytrids, 57 

Anisolpidiaceae, 69 

Anisoplidium, 69 

Anisolpidium Ectocarpii, 70 

Anisonema, 109 

Anisonema truncatun, 108 

Anisonemida, 105, 108 

Anisonemidae, 108 

Anisonemina, 108 

Anomalinidae, 187 

Anopheles, 213 

Anoplophryida, 230 

Anoplophryinca, 229, 230 

Anthophysis, 59 

Anucleobionta, 6, 12 

Ape, 213 

Aphanizomenon, 35 

Aphanomycopsis, 81 

Aphrothoraca, 190, 193 

Aphrothoracida, 190 

Aplanosporeao, 86 

Aplosporidics, 218 

Apodachlya, 79 

Apodachlyclla, 79 

Apodinidae, 102 

Apodinium, 102 

Appcndiculatae, 73 



Apple, 139, 148 
Araceae, 67 
Arachnula, 191 
Araiospora, 79 
Arcella, 205 
Arcellidae, 205 
ArcelHna, 205 
Archaelagena, 186 
Archaias, 184, 185 
Archangiacea, 28 
Archangiaceae, 28 
Archangium, 28 
Archegregarina, 215 
Archephyta, 17 
Archezoa (of Haeckel), 17 
Archezoa (of Perty}, 223 
Archi-Monothalamia, 183 
Archimycetae, 110 
Archimycetes, 110, 111 
Archiplastidea, 18, 29 
Archiplastideae, 30 
Arcyria, 176 
Arcyriacea, 174, 176 
Arcyriaceae, 176 
Arcyriidae, 176 

Arthropods, 211, 212, 222 

Arthrospira, 35 

Asclepiadaceae, 161 

Ascobolacea, 135 

Ascochyta, 141 

Ascocorticium, 137, 145 

Ascocyclus, 88 

Ascoidea, 130 

Ascoidea rubescens, 127 

Ascoideaceae, 130 

Ascomvcetae, 125 

Ascomycetes, 120, 125, 140, 142, 145 

Ascomyceten, 125 

Ascosporeae, 125 

Askcleta, 193 

Aspergillus, 130, 131 

Aspergilliales, 130 

Aspidisca, 232 

Asplrotricha, 229 

Aspirotrichaceae, 229 

Astasia, 96, 107 

Astasiaceae, 107 

Astasiaca, 96, 105, 107 

Astasiidae, 107 

Astasiina, 107 

Asterigerina, 187 

Asterigerinida, 187 

Asterigerinidae, 187 

Asterocyclina, 188 

Asterocystis, 43 

Asterophlyctis, 1 1 7 

Astoma, 94, 96, 105 

Astoinaticae, 74 

Astomina, 230 

Astracanthida, 199 

Astracanthidae, 199 

Astrodisculus, 193 

Astrolophida, 197 
Astrolophus, 197 
Astrorhiza, 183 
Astrorhizida, 183 
Astrorhizidaceae, 183 
Astrorhizidae, 183 
Astrorhizidea, 183 
Astrorhizina, 183 
Ataxophragmidae, 186 
Ataxophragmidea, 186 
Ataxophragmium, 186 
Athene noctua, 162 
Aulacantha, 199 
Aulacanthida, 199 
Aulacanthidae, 199 
Aulactinium, 199 
Aulosphaera, 199 
Aulosphaerida, 199 
Auricularia, l46 
Auricularia Auricula, 146 
Auriculariacea, 146, 148 
Auriculariaceae, 146 
Auriculariales, 146 
Auriculariineae, 145, 146 
Auricularineae, 146 
Autobasidiomycetes, 146 
Aves, 6 

Axonoblasteae, 51 
Azoosporidae, 191 
Azoosporidca, 191 
Azoosporidia, 190 
Azotobacter, 14 
Azotobacter Chroococcum, 23 

Azotobacteriacca, 19, 23 
Azotobacteriaceae, 23 
Babesia, 214 

Babesia bigemina, 206, 212, 214 
Babesiida, 211, 214 
Babesiidae, 214 
Bacillacea, 19, 21 
Bacillacei, 21 
Bacillaria, 69, 75 
Bacillariacea, 1 1, 55, 65, 69, 72 
Eacillariaceae, 71 
Bacillariales, 53, 71 
Bacillarieae, 71 
Bacillarioideae, 71 
Bacillariophyceae, 71 
Bacillariophyta, 71 
Bacillus, 21 
Bacillus alvei, 21 
Bacillus Amylobacter, 21 
Bacillus anthracis, 21 
Bacillus, colon, 22 
Bacillus, gas, 22 
Bacillus Radicicola, 23 
Bacillus, Shiga, 22 
Bacillus subtilis, 18, 21 
Bacteria, 2, 3, 4, 6, 7, 12, 13, 14, 17, 18, 
30, 38, 118, 119, 189, 222, 224, 231 


The Classification of Lower Organisms 

Bacteriaceae, 21 

Bacteriophyta, 17 

Bacteroides, 22 

Badhamia, 177 

Balantidium, 230 

Balantidium coli, 230 

Bangia, 43 

Bangia fuscopurpurea, 43 

Bangiacea, 41 

Bangiaceae, 41, 43 

Bangialea, 40, 41, 52 

Bangiales, 41 

Bangieae, 41 

Bangiineae, 41 

Bangioideae, 41 

Barbulanympha, 169 

Barley, 6 

Barrouxia, 210 

Bartonella bacilliformis, 21, 214 

Bartonellaceae, 20 

Bartramia, 219 

Bartramiidae, 218 

Basidiobolacea, 125 

Basidiobolaceae, 125 

Basidiobolus, 119, 121 

Basidiobolus ranarum, 125 

Basidiomycetae, 142 

Basidiomyceten, 142 

Basidiomycetes, 121, 127, 128, 141, 142, 

Basidiosporeae, 142 

Bathysiphon, 183 

Batrachospermaceae, 47 

Batrachospermum, 47 

Bdellospora, 124 

Beetles, 177, 215, 217 

Beggiatoa, 24, 30, 31, 32, 35 

Beggiatoacea, 34, 35 

Beggiatoaceae, 35 

Bicoecaceae, 67 

Bicoecidea, 67 

Bicoekida, 67 

Bicosoeca, 67 

Biddulphia, 74 

Biddulphiaceae, 74 

Biddulphica, 74 

Biddulphicac, 74 

Biflagcllatae, 76 

Bikoecidae, 67 

Bikoccina, 67 

Birds, 6, 210. 212, 213 

Bitunicatae, 129 

Blakeslcca, 124 

Blastocaulis, 26, 27 

Blastocladia, 112, 113 

Blastocladiarca, 110, 112 

Blastocladiaceae, 112 

Blastocladialcs, 1 1 1 

Blastocladiclla, 112, 113 

Blastocladiclla cystogena, 115 

Blastocladiineae, 111 

Blastodcrma, 130 

Blastodinida, 100, 102 

Blastodinidae, 102 

Blastodinides, 102 

Blastodinium, 102 

Blastosporaceae, 44 

Blepharisma, 230 

Blue grass, 148 

Blue-green algae, see Algae, Blue-green 

Bodo, 159, 160, 199, 209, 212 

Bodo edax, 161 

Bodo Lacertae, 159 

Bodonaceae, 159 

Bodonidac, 159 

Bodonidca, 158 

Bodonina, 159 

Boletus, 151 

Bolivina, 188 

Borelis, 185 

Borrelia, 29 

Borrelia recurrentis, 28, 29 

Borrelia Vincenti, 29 

Botrida, 198 

Botrydiaceae, 67 

Botrydiales, 63 

Botrydiopsis, 66 

Botrydium, 65, 67 

Botryococcacea, 65, 66 

Botryococcaceae, 66 

Botryococcus, 66 

Botryoglossum, 52 

Botryoidca, 198 

Botrytis, 140, 142 

Bovista, 155 

Braadrudosphaeridae, 60 

Brachycystida, 217 

Brefcldia, 175 

Brefeldiaceae. 175 

Brefeldiidae, 175 

Brehmiella, 59 

Brehmiella chrysohydra, 54 

Brown algae, see Algae, Brown 

Brucella, 22 

Bulgariacea, 135 

Bulimina, 188 

Buliminida, 188 

Buliniinina, 188 

Bumilleria, 66, 73 

Bursaria, 230 

Bursariidae, 230 

Bursarina, 230 

Cabbage, 1 78 
Calcaroae, 1 71 
Calcarina, 187 
Calcarinidae, 187 
Calciconus, 60 
Galciconus vitrcus, 56 
Calrisolcnia. 60 
Calcisolonidae, 60 
Callocolax, 50 
Callophyllis, 50 



Calonectria, 142 
Calonema, 177 
Calonemeae, 171 
Calonympha, 168 
Calonymphida, 166, 167, 168 
Calonymphidae, 168 
Calothrix, 36 
Calvatia, 155 
Calyptosphaera, 60 
Calyptosphaera insignis, 56 
Camerina, 188 
Camerinidae, 188 
Camptonema, 193 
Camptonematidae, 193 
Campuscus, 191 
Candida, 142 
Cannobotryida, 198 
Cannopilus, 63 
Cannosphaerida, 199 
Cannosphaeridae, 199 
Cantharellales, 150 
Carageen, 49 
Carboxidomonas, 24 
Carchesium, 233 
Carcheslum polypinum, 225 
Carpomitra, 88 
Carpomycetae, 119 
Carpophyceae, 40 
Carposporeen, 128 
Caryococcus, 21 
Caryospora, 210 
Caryotropha, 211 
Cassidulina, 188 
Cassidulinida, 188 
Cassidulinidae, 188 

Castanellida, 200 

Castanellidae, 200 

Castanidium, 200 

Cat, 6, 210 

Catenariopsis, 69 

Catenochytridium, 118 

Cattle, 206, 214 

Caulleryella, 215 

CauUeryellidae, 215 

Caulobacter, 26, 27 

Caulobacter vibrioides, 26 

Caulobacteriacea, 27 

Caulobacteriaceae, 27 

Caulobacterialea, 18, 25, 26 

Caulobacteriales, 25 

Cayeuxina, 186 

Cellulomonas, 22 

Cenolarcus, 195 

Centipedes, 207, 210, 211 

Centricae, 73, 74 

Cepedia, 229 

Cephalina, 217 

Cephalopodes, 182 

Cephalothamnium, 59 

Cephalothamnium Cyclopum, 54 

Cephalotrichinae, 18 

Ceramiales, 51 

Ceramiea, 51, 52 
Ceramieae, 51 
Ceratiidae, 103 
Ceratiomyxa, 177, 221 
Ceratiomyxa fruticulosa, 177, 178 
Ceratiomyxacea, 177 
Ceratiomyxaceae, 177 
Ceratium (dinoflagellate), 103 
Ceratium ( myxomycete ) , 177 
Ceratium Hirundinella, 103 
Ceratomyxa, 221 
Ceratomyxidae, 221 
Ceratophyllus fasciatus, 160 
Ceratospora, 216 
Cercobodo, 159 
Cercobodonidae, 159 
Cercomonadida, 159 
Cercomonadidae, 159 
Cercomonadinea, 158 
Cercomonas, 159, 161 
Cercomonas Davainei, 165 
Cercomonas Hominis, 165 
Cercomonas longicauda, 160 
Cercomonas obliqua, 165 
Cercospora, 138, 139, 142 
Chaenia, 229 
Chaetangieae, 47 
Chaetoceraceae, 74 
Chaetoceros, 74 
Chaetocladiaceae, 124 
Chaetocladium, 123, 124 
Chaetoproteida, 159, 163 
Chaetoproteidae, 163 

Chaetoproteus, 158, 160, 163, 202 

Chaidae, 201 

Chaidea, 201 

Chalarothoraca, 190, 193 

Chalarothoracida, 190 

Challengerida, 200 

Challengeridae, 200 

Challengeron, 200 

Chamaesiphon, 35, 36 

Chamaesiphon incrustans, 32 

Chamaesiphonacea, 34, 35 

Chamaesiphonaceae, 33, 35 

Champia, 51 

Champiea, 51 

Champieae, 51 

Chantransia, 47 

Chantransiaceae, 47 

Chaos Protheus, 200, 201, 202 

Chaosidae, 201 

Chapmania, 187 

Chapmaniida, 187 

Chapmaniidae, 187 

Characiopsis, 66 

Characiopsis gibba, 64 

Chestnut, 139 

Chiastolida, 197 

Chiastolus, 197 

Chicken, 210 

Chilodon, 230 


The Classification of Lower Organisms 

Chilodon uncinatus, 225 
Chilodontida, 230 
Chilomastigidae, 165 
Chilomastix, 165 
Chilomastix davainei, 165 
Chilomastix Hominis, 165, 237 
Chilomastix Mesnili, 165 
Chilomonadaceac, 98 
Chilomonas, 94, 109 
Chilomonas Paramaecium, 97 
Chilostomella, 187 
Chilostomellida, 187 
Chilostomellidae, 187 
Chlamydodon, 230 
Chlamydodonta, 230 
Chlamydodontida, 230 
Chlamydodontidae, 230 
Chlamydomonas, 61, 111 
Chlamydomyxa, 191 
Chlamydomyxidea, 190 
Chlamydophora, 190, 193 
Chlamydophorida, 190 
Chlamydothrix ochracea, 32, 36 
Chlamydotrichacea, 34 
Chlamydotrichaceae, 36 
Chlamydozoaceae, 20 
Chloramoeba, 66 
Chloramoeba heteromorpha, 64 
Chloramoebacca, 65, 66 
Chloramoebaceae, 66 
Chloramoebidac, 66 
Chlorarachnidae, 66 
Chlorobacteriaceae, 31 
Chlorobacterium, 33 
Chlorobiacea, 31, 237 
Chlorobium, 31 
Chlorobotrydiaceae, 66 
Chlorochromonas, 66 
Chlorochytridion, 1 1 1 
Chloromonadaceae, 109 
Chloromonadales, 63, 105 
Chloromonadida, 105 
Chloromonadidae, 109 
Chloromonadina, 63, 96, 105 
Chloromonadinae, 94, 105 
Chloromonadineae, 105 
Chloromonads, 94 
Chloromyxea, 221 
Chloromyxees, 221 
Chloromyxida, 221 
(^hloromyxidac, 221 
Chloromyxum, 221 
Chlorosaccacca, 65 
Chlorosaccaceac, 65 
Chlorosaccus, 55, 65, 66 
Chlorosaccus fluidus, 64 
Chlorotheciacea, 65, 66 
Chlorothcciaceae, 66 
Choancphoraccac, 124 
Choano-Flagellata, 67 
Choanocystidac, 194 
Choanocystis, 194, 216 

Choanoflagellata, 57, 61, 67, 68 
Choanoflagcllates, 57, 158 
Choanosporidae, 216 
Chondria, 52 
Chondrieae, 51 
Chondrioderma, 177 
Chondrococcus, 28 
Chondromyces, 28 
Chondromyces aurantiacus, 26 
Chondromyces crocatus, 26 
Chondrus, 51 
Chondrus crispus, 49 
Chonotricha, 230, 231, 237 
Chonotrichida, 230 
Chordariacea, 88 
Chordariaceae, 87 
Chordariales, 87 
Chordarieae, 87 
Chromatiacea, 31, 237 
Chromatiaceae, 31 
Chromatium, 31 
Chromobacterium, 22 
Chromomonas, 98 
Chromulina, 61, 62 
Chromulina Pascheri, 56 
Chromulinaceae, 62 
Chromulinales, 61 
Chromulinidae, 62 
Chroococcacea, 33 
Chroococcaceae, 33 
Chroococcales, 33 
Chroococcus, 32, 33 
Chrysamoeba, 63 
Chrysamoebida, 62, 63 
Chrysamoebidae, 63 
Chrysapsis, 62 
Chrysarachniaceae, 63 
Chrysarachnion, 63 
Chrysidella, 98 
Chrysocapsa, 59 
Chrysocapsa paludosa, 54 
Chrysocapsacea, 58, 59 
Chrysocapsaceae, 59 
Chrysocapsales, 61 
Chrysocapsidae, 59 
Chrysocapsina, 61 
Chrysocapsinae, 61 
Chrysocapsineae, 55, 61 
Chrysochromulina, 58 
Chrysococcus, 62 
Chrysocrinus, 63 
Chrysodcndron, 59 
Chrysomonadaceae, 59 
Chrysomonadales, 61 
Chrysomonadida, 61 
Chrysoinonadidao, 62 
Chrysomonadina, 59, 61, 62 
Chrysomonadinae, 61 
Chrysomonadinca, 57 
Chrysoinonadincac, 55, 57, 61 
Chrysonionads, 53, 83 
Chrysomonas, 62 



Chrysophaeum, 109 

Chrysophyceae, 53, 55, 95 

Chrysophycophyta, 53 

Chrysophyta, 53 

Chrysopyxis, 60 

Chrysosphaera, 62 

Chrysosphaeracea, 61, 62 

Chrysosphaeraceae, 62 

Chrysosphaerales, 61 

Chrysosphaerella, 62 

Chrysosphaerineae, 55, 61 

Chrysospora, 62 

Chrysothylakion, 63 

Chrysotrichaceae, 60 

Chrysotrichales, 61 

Chrysotrichineae, 55, 61 

Chytridiacea, 117, 118 

Chytridiaceae, 110, 118 

Chytridiales, 113 

Chytridieae, 110 

Chytridieen, 110, 118 

Chytridiineae, 110 

Chytridinae, 110 

Chytridinea, 111, 113, 116 

Chytridineae, 110, 113 

Chytridium, 69, 110, 113, 118 

Chytridium Olla, 110 

Chytrids, 76, 110, 111, 119, 121, 125, 130, 

Chytriodinium, 102 
Cienkowskiaccae, 177 
Ciliata, 223, 228, 233 
Ciliatea, 228 
Cilio-flasrellata, 94 
Cilioflagellata, 96, 102 
Giliophora, 39, 223, 237 
Ciliophryidae, 191 
Ciliophrys, 193 
Circoporida, 200 
Circoporidae, 200 
Circoporus, 200 
Cladochytriacea, 115, 117 
Cladochytriaceae, 117 
Cladochytrium, 110, 117 
Cladococcida, 195 
Cladococcus, 195 
Cladopyxida, 103 
Cladosporium, 142 
Cladothrix dichotoma, 33 
Clastoderma, 175 
Clathracea, 155 
Clathraceae, 155 
Clathrochloris, 31 
Clathrulina, 194 
Clathrulinida, 191, 194 
Clathrulinidae, 194 
Claudea, 51 
Clavaria, 151 
Clavariacea, 151 
Clavariaceae, 151 
Clavariei, 151 
Clavati, 150 

Claviceps purpurea, 139 
Clonothrix fusca, 32, 36 
Closterium, 125 
Clostridium, 21 
Clostridium botulinum, 21 
Clostridium butyricum, 21 
Clostridium Pastorianum, 21 
Clostridium septicum, 21 
Clostridium tetani, 21 
Cnemidospora, 21 7 
Cnidosporidea, 219 
Cnidosporidia, 219, 220 
Coccaceac, 20 
Coccidia, 207, 210 
Coccidians, 260, 209, 210, 212, 215 
Coccididae, 210 
Coccidiidea, 210 
Coccidiomorpha, 207, 210 
Coccidium, 210 
Coccidium Schubergi, 207 
Coccogonales, 33 
Coccogonea, 31, 32, 33 
Coccogoneae, 33 
Coccolithaceae, 60 
Coccolithidae, 60 
Cocclithina, 60 
Coccolithophora, 60 
Coccolithophoridae, 55, 60 
Coccolithus, 60 
Coccomyces, 134 
Coccomyxa, 221 
Coccomyxida, 221 
Coccomyxidae, 221 
Cocconeidaceae, 76 
Cocconeis, 72, 73, 76 
Cocconemaceae, 75 
Cocconemidae, 222 
Coccosphaera, 60 
Coccospora Slavinae, 222 
Coccosporida, 222 
Coccosporidae, 222 
Coccus, 20 
Cochliodinium, 101 
Cochliopodiidae, 202 
Cochliopodium, 202 
Cochlonema, 124 
Cockroach, 169,217,219 
Codonoecina, 67 
Codonosiga, 67 
Codonosigidae, 67 
Codosiga, 67 
Coeloblastca, 46 
Coeloblasteae, 51 
Coclodendrida, 200 
Coelodendrum, 199, 200 
Coelomonadina, 105, 109 
Coelosphaerium, 33 
Coelosporidiidae, 218 
Coelosporidium, 219 
Coenenia, 203 
Coffee, 148 
Colaciacea, 105 


The Classification of Lower Organisms 

Colaciaceae, 105 
Colaciidae, 105 
Colacium, 105 
Colacium Arbuscula, 106 
Coleosporiacea, 148 
Coleosporiaceae, 148 
Coleosporium, 143 
Coleosporium Vernoniae, 143 
Colepina, 229 
Coleps, 229 

Colletotrichum, 139, 140 
Collida, 195 
Collodaria, 194 
Colloderma, 177 
Collodermataceae, 177 
CoUosphaera, 195 
Collosphaera Huxleyi, 196 
Collosphaerida, 195 
Collozoida, 195 
Colpidium campylum, 227 
Colpoda, 229 
Colpodaea, 229 
Colpodella, 189 
Colpodidae, 229 
Columniferae, 171 
Comatricha, 175 
Completoria, 125 
Compsopogon, 44 
Compsopogonacea, 41, 44 
Compsopogonaceae, 44 
Concharida, 200 
Concharidae, 200 
Conchulina, 205 
Conferva, 66 
Confervaceae, 66 
Confervales, 63 
Confervoidea, 63 
Conger niger, 161 
Conidiobolus, 125 
Coniferinae, 9 
Conifers, 148 
Coniomycetes, 140 
Conjugatae, 117 
Conradiella, 62 
Coprinus, 143, 152 
Coprinus atramentarias, 153 
Copromonas subtilis, 108 
Cora, 151 
Corallinaceae, 50 
Corallinea, 50 
Corallineae, 50 
Cordyceps, 139 
Coreocolax, 50 
Corethron, 74 
Cormobionta, 6 
Cornuspira, 185 
Coronympha, 168 
Corticiiun, 151 
Corynebacteriacea, 19, 20 
Coryncbacteriaceae, 20 
Corynebacteriidae, 20 
Coryncbactcrium, 20, 21 

Corynebacterium diphtheriae, 20 
Coryneum, 141 
Coryneum Beijerinckii, 141 
Coscinodiscaceae, 74 
Coscinodiscea, 74 
Coscinodiscus, 74 
Costia, 165 
Costiidae, 165 
Cothurnia, 233 
Councilmania, 203 
Crab, 218 
Craigia, 163 

Craspedomonadaceae, 67 
Craspedomonadina, 67 
Craspedotella, 102 
Craterellus, 151 
Craterium, 177 
Crenothrix polyspora, 32, 36 
Crenotrichacca, 35, 36 
Crenotrichaceae, 36 
Cribraria, 175 
Cribrariacea, 173, 175 
Cribrariaceae, 171, 175 
Cribrariales, 171, 173 
Cribrariidae, 175 
Cribrospira, 186 
Cristellaria, 187 
Cristispira, 29 
Cristispira Veneris, 26 
Crithidia, 162 
Cromodromys, 199 
Cronartiacea, 148 
Cronartiaceae, 148 
Cronartium, 148 
Cronartium ribicola, 148 
Cryptobia, 160, 161,209,212 
Cryptobiidae, 161 
Cryptocalpis, 198 
Cryptocapsales, 97 
Cryptocapsineae, 95 
Cryptocercus, 166, 169, 170 
Cryptochrysis, 98 
Cryptococcacea, 97, 98 
Cryptococcaceae, 98 
Cryptococcalcs, 96, 97 
Cryptococcineae, 95 
Cryptococcus, 98, 130 
Cryptocystes, 219, 222 
Cryptomonadaceae, 98 
Cryptomonadalca, 96 
Cryptomonadalcs, 96 
Cryptomonadida, 97 
Cryptomonadidae, 98 
Cryptomonadina, 96, 97, 98 
Cryptonionadinae, 96 
Cryptonionadincae, 95, 96 
Crvptomonads, 94, 194 
Cryptomonas, 97,98, 199 
Cryptoncnicac, 50 
Cryptonemiales, 50 
Cryptoncniinae, 50 
Cryptophyceae, 94, 96 



Cryptospermea, 46, 47 
Cryptospermeae, 47 
Ctenomyces, 131 
Ctenostomata, 230, 231 
Ctenostomida, 231 
Ctenostomina, 231 
Cumagloia, 47 
Cuneolina, 186 
Cunninghamella, 124 
Cup fungi, 134 
Cupulata, 129, 134, 137, 145 
Cupulati, 134 
Currants, 148 
Cutleria, 88 
Cutlcriacea, 88 
Cutlerialea, 85, 88 
Cutleriales, 88 
Cyanomonas, 98 
Cyanophyceae, 29 
Cyanophyta, 17, 30 
Cyathoxnonas, 97, 98 
Cyathus, 155 
Cyclammina, 186 
Gyclidina, 230 
Cyclidium, 230 
Cycloclypeidae, 188 
Cycloclypeina, 188 
Cycloclypeus Carpenter!, 188 
Cyclodinidae, 229 
Cyclonexis, 59 
Cyclonympha, 171 
Cyclonymphidae, 169 
Cycloposthium, 231 
Gyclosiphon, 188 
Cyclosporales, 91 
Cyclosporeae, 82, 91 
Cyclotella, 72, 73, 74 
Cylindrospermum, 35 
Cylindrospermum majus, 32 
Cylindrosporium Pruni, 134 
Cymbalopora, 180, 182, 187 
Cymbella, 72, 73, 75 
Cymbellea, 75 
Cymbelleae, 75 
Cyphoderia, 191 
Cyrtellaria, 198 
Cyrtida, 198 
Cyrtoidea, 198 
Cyrtophora, 62, 63 
Cystidium, 198 
Cystobasidium, 147 
Cystobasidium sebaceum, 145 
Cystochytrium, 69 
Cystoflagellata, 94, 96, 99 
Cytophaga Hutchinsonii, 26, 28 
Cytophagacea, 28 
Cytophagaceae, 28 
Cytosporidia, 207 
Cyttariacea, 135 

Dacryomitra, 150 
Dacryomyces, 150 
Dacryomycetacea, 150 
Dacryomycetaceae, 150 
Dacryomycetalea, 146, 150 
Dacryomycetales, 150 
Dacryomycetineae, 150 
Dactylophorida, 218 
Dactylophoridae, 218 
Dactylophorus, 218 
Dactylosphaerium, 202 
Daedalea, 151 
Daldinia, 139 
Dallingeria, 58 
Dasyea, 51 
Daucina, 188 
Deer, 214 
Delacroixia, 125 
Delesseria, 51 
Delesseria sinuosa, 49 
Delesseriea, 51 
Dematiaceae, 142 
Dematiea, 142 
Dematieae, 142 
Dematiei, 141 
Dendrocometes, 236 
Dendrocometida, 236 
Dendrocometidae, 236 
Dendrocometina, 236 
Dendromonadina, 59 
Dendromonas, 59 
Dendromonas virgaria, 54 
Dendrosoma, 235 
Dendrosomatidae, 235 
Dendrosomida, 235 
Dendrosomidae, 235 
Dendrosomina, 235 
Dentilina, 184 
Derepyxis, 60 
Dermateacea, 135 
Dermatocarpa, 146, 152 
Dermatocarpi, 152, 171 
Dermocarpa, 36 
Dermocarpa protea, 32 
Dermocentor, 20 
Desmarestia, 88, 89 
Desmarestiacea, 88 
Desmarestales, 87 
Desmobacteriales, 33 
Desmocapsa, 99 
Desmocapsales, 98, 99 
Desmocapsineae, 95, 99 
Desmokontae, 94, 98, 99 
Desmomastix, 99 
Desmomonadales, 98, 99 
Desmomonadineae, 95, 99 
Desmothoraca, 190, 194 
Desmothoracida, 190 
Desmotrichum, 88 
Deuteromycetes, 140 
Deutschlandiaceae, 60 
Devescovina, 167 


The Classification of Lower Organisms 

Devescovinida, 167 
Devescovinidae, 167 
Devescovininae, 167 
Diachea, 175 
Dianema, 176 
Dianemaceae, 176 
Diaporthe, 139 
Diatoma, 75 
Diatomaceae, 69, 75 
Diatomea, 53, 69, 71, 74 
Diatomeae, 53, 69, 71, 74 
Diatoms, 53, 71,83, 117, 118 
Diatrype, 139 
Dictydiaethaliaceae, 175 
Dictydiaethaliidae, 1 75 
Dictydiaethalium, 175 
Dictydium, 175 
Dictyocha, 63 
Dictyocha Fibula, 56 
Dictyochaceae, 62 
Dictyochidae, 62 
Dictyoconoides, 187 
Dictyoconus, 186, 198 
Dictyophora, 155 
Dictyosiphonales, 89, 91 
Dictyosteliaceae, 203 
Dictyosteliaceen, 203 
Dictyostelidae, 203 
Dictyostelium, 203 
Dictyostclium discoideum, 204 
Dictyostelium mucoroides, 204 
Dictyota, 87 
Dictyotacea, 87 
Dictyotaceae, 86, 87 
Dictyotales, 82, 86 
Dictyotea, 85, 86 
Dictyotcae, 82, 86 
Dictyuchus, 78, 79 
Didesmis, 231 
Didiniidae, 229 
Didinium, 229, 235' 
Didinium nasutum, 225 
Didymiacea, 175, 177 
Didymiaceae, 177 
Didymidae, 177 
Didymiidae, 177 
Didymium, 177 
Didymohelix ferruglnea, 27 
Didymophyes, 218 
Didymophyida, 218 
Didymophyidac, 218 
Difflugia, 201, 205 
DIfflugiida, 205 
Difflugiidae, 205 
Dilcptus, 230 
Dimastigamocba, 159 
Dimorpha, 193 
Dimychota, 17 

Dinamocba (dinoflagellate), 101 
Dinamocba (amoeba), 16, 202 
Dinamocbidina, 100, 101, 237 
Dinamocbidium varians, 101, 104 

Dinastridium, 100 
Dinenympha, 166 
Dinenymphida, 165, 166 
Dinenymphidae, 166 
Dinifera, 102 
Diniferidea, 103 
Dinobryaceae, 60 
Dinobryina, 58, 60 
Dinobryon, 58, 60 
Dinocapsaceae, 100 
Dinocapsales, 99, 100 
Dinocapsina, 99 
Dinocapsineae, 95, 99 
Dinococcales, 99, 100 
Dinococcina, 99 
Dinococcineae, 96, 100 
Dinoclonium, 100 
Dinocloniaceae, 100 
Dinoflagellata, 94, 102 
Dinoflagellatae, 94, 95 
Dinoflagellates, 94, 199 
Dinoflagellida, 103 
Dinophyceae, 94, 103 
Dinophysida, 103 
Dinophysis, 103 
Dinothrix, 100 
Dinotrichales, 99, 100 
Dinotrichineae, 96, 99 
Dioxys, 66 
Dioxys Incus, 64 
Diplococcus, 20 
Diplococcus pneumoniae, 20 
Diploconida, 197 
Diploconus, 197 
Diplocystida, 216 
Diplocystidae, 216 
Diplocystis, 216 
Diplodia, 141 
Diplodinium, 224, 231 
Diplomita, 60 
Diplophlyctis, 117 
Diplophysalis, 191 
Diplophysalis stagnalis, 192 
Dipodascus, 130 
Dipodascus albidus, 132 
Discellacea, 141 
Discellaceae, 141 
Dischizae, 21 5 
Discida, 195 
Disciformia, 73 
Discoasteridac, 60 
Dificoidca, 195 
Discolichenes, 134 
Discomycetes, 133, 134 
Discophrya, 235 
Discophryida, 235 
Discophryidae, 235 
Discorbis, 180, 182 
Discorbis mcditerrancnsis, 182 
Discorbis orbicularis, 182 
Discosphacra, 60 
Disporees, 209 



Distephanus, 63 

Distephanus Speculum, 56 

Distigma, 107 

Distomata, 163 

Distomataceae, 166 

Distomatinales, 163 

Distomatineae, 163 

Ditripodiidae, 62 

Doassansia, 149 

Dobellia binucleata, 210 

Dobeliida, 210 

Dobelliidae, 210 

Do?, 210 

Dolichocystida, 209, 214 

Doliocystida, 216 

Doliocystidae, 216 

Doliocystis, 216 

Dorataspida, 197 

Dorataspis, 197 

Dorataspis costata, 196 

Dothideaceae, 137 

Dothideales, 137, 138, 139, 140, 141 

Drepanidium, 211 

Duboscqia, 222 

Dudresnaya purpurifera, 49 

Dumontieae, 50 

Earth star, 155 
Earthworm, 215, 216 
Eberthella, 22 
Eberthella typhi, 22 
Ebriaceae, 55, 62 
Ebriidae, 62 
Ebriopsidae, 62 
Echinocystida, 189 
Echinoderms, 216 
Echinosteliaceae, 175 
Echinostelium, 175 
Ectocarpales, 86 
Ectocarpea, 86 
Ectocarpeae, 86 
Ectocarpineae, 86 
Ectocarpus, 70, 83, 86, 87 
Ectocarpus Mitchelliae, 204 
Ectocarpus siliculosus, 83 
Ectosporeae, 177 
Ectrogella, 81 
Ectrogellacea, 81 
Ectrogellaceae, 81 
Eel, 161 

Egregia Menziesii, 90, 91 
Eimeria, 210 
Eimerida, 210 
Eimeridae, 210 
Eimeridea, 210 
Eimcriidea, 210 
Eimeriinea, 210 
Eimerioidae, 210 
Elaeorhanis, 193 
Elaphomyces, 131 
Ellipsoidina, 188 

Elphidium, 186, 187 
Elphidium crispum, 181 
Elvella, 135 
Empusa, 125 
Enchelia, 229 
Enchelina, 229 
Enchelis, 229 
Enchelyidae, 229 
Endamoeba, 202, 203 
Endamoeba disparita, 202 
Endamoeba histolytica, 202 
Endamoebida, 201, 202 
Endamoebidae, 202 
Endocochlus, 124 
Endogonacea, 123, 124 
Endogonaceae, 124 
Endogone, 123, 124 
Endogonei, 124 
Endoiimax, 203 
Endomyces, 130 
Endomycetacea, 130 
Endomycetaceae, 130 
Endomycetalea, 129 
Endomycetalcs, 129 
Endosporea, 171 
Endosporeae, 171 
Endosporinei, 171 
Endothia parasitica, 139 
Endothyra, 186 
Endothyridae, 186 
Endothyrina, 186 
Enerthenema, 175 
Enerthenemaceae, 175 
Enerthenemea, 174, 175 
Entamoeba, 202, 203 
Entamoeba coli, 202 
Entamoeba dystenteriae, 202 
Entamoeba gingivalis, 202 
Enteridiea, 171 
Enteridieae, 171 
Enterobacteriaceae, 21 
Entodiniomorpha, 230, 231 
Entodiniomorphina, 231 
Entodinium, 231 
Entomophthora, 125 
Entomophthoracea, 124 
Entomophthoraceae, 124 
Entomophthorales, 124 
Entomophthorinea, 121, 124 
Entomophthorineae, 124 
Entophlycis, 113, 117 
Entophysalidales, 33 
Entosiphon sulcatum, 108 
Eocronartium, 143, 147 
Eocronartium muscicola, 145 
Eouvigerina, 188 
Ephelota, 236 
Ephelotida, 236 
Ephelotidae, 236 
Ephelotina, 236 
Ephemera vulgata, 222 
Epiblasteae, 50 


The Classification of Lower Organisms 

Epichrysis, 56, 62 

Epiclintes, 233 

Epidinium, 231 

Epipyxis, 60 

Epipyxis utriculus, 54 

Epistylis, 233 

Eremascus, 130 

Eremascus albidus, 127 

Eremospermeae, 77 

Erica, 9 

Ericae, 9 

Erysiphe, 127, 132, 133 

Erysiphe graminis, 132 

Erysiphea, 133 

Erysipheae, 133 

Erythrocladia, 44 

Erythropsis, 101 

Erythrotrichia, 44 

Erythrotrichia carnea, 44 

Erythrotrichiaceae, 44 

Erwina, 22 

Erwinia amylovora, 22 

Escherichia coli, 14, 15, 22 

Ethmosphaerida, 195 

Euactinomyxidae, 222 

Euasci, 130 

Eubacteria, 18, 25 

Eubacteriales, 18 

Eubasidii, 145 

Euchrysomonadina, 61 

Euchrysomonadinae, 61 

Eucomonympha, 169 

Eucyrtidina, 198 

Eucyrtidium, 198 

Eucyrtidium carinatum, 196 

Eudesme, 88 

Euflorideae, 44 

Euglena, 38, 94, 107, 116, 117, 125 

Euglena acus, 106 

Euglena Spirogyra, 106, 107 

Euglena viridis, 106 

Euglenaceae, 105 

Englenales, 105 

Euglenamorpha, 105 

Euglenida, 105 

Euglenids, 94, 106 

Euglenina, 105 

Euglcninae, 94, 105 

Euglenineae, 96, 105 

Euglenocapsineae, 96 

Euglcnoidina, 96, 105 

Euglenophycophyta, 94 

Euglenophyta, 94 

Euglypha, 191 

Euglyphida, 191 

Euglyphidae, 191 

Eugregarinaria, 217 

Eugregarinida, 217 

Eumycetes, 119 

Eumycctozoina, 171 

Eumycophyta, 119 

Eunotia, 75 

Eunotiaceae, 75 
Eunotiea, 75 
Eunotieae, 75 
Euphorbiaceae, 161 
Euplotes, 227, 232, 233, 234 
Euplotes Patella, 225, 226, 232 
Eupodiscales, 73 
Eurotium, 131 
Eurychasma, 81 
Eur)'chasmidium, 81 
Eurysporea, 221 
Eutreptia, 105 
Excipula, 141 
Excipulaceae, 141 
Exidia, 143 
Exoascalea, 129, 137 
Exoascales, 137 
Exoascus, 137 
Exobasidiacea, 151 
Exobasidiaceae, 151 
Exobasidiales, 1 50 
Exobasidiineae, 150 
Exobasidium, 151 
Exosporea, 171, 177 
Exosporeae, 177 
Exosporinei, 177 
Exuviaella, 99 

Fasciolites, 185 

Fauchea, 51 

Faucheocolax, 51 

Felis Catus, 6 

Ferns, 125, 148 

Filicineae, 1 

Fisherinidae, 185 

Fishes, 165, 210, 211, 219. 220, 222 

Flabellina, 184, 187 

Flagellata, 6, 55, 94, 96, 105 

Flagellatae, 94 

Flagellates, 10, 53, 55, 76, 94, 118 

Flagellato-Eustomata, 105 

Flagellato-Pantostomata, 158 

Flatworms, 216 

Flavobacterium, 22 

Flea, 160 

Flexostylida, 185 

Floridea, 47, 50,51 

Florideae, 6, 40, 44, 51 

Floridees, 40, 51 

Floridineae, 44 

Flowers of tan, 177 

Fly, 213 

Foaina, 167 

Folliculina, 230 

Fomcs, 151 

Foraminifera, 179, 182, 183, 185 

Foraminiferes, 179, 182 

Foraminiferida, 179 

Forficule, 217 

Fragilaria, 75 

Fragilariaceae, 75 



Fragilariea, 75 

Fragilarieae, 75 

Frogs, 125, 210, 211 

Frondicularia, 187 

Fucaceae, 91 

Fucales, 91 

Fucea, 91 

Fuceae, 91 

Fucineae, 91 

Fucacees, 82 

Fucoidea, 83, 86, 91 

Fucoideae, 53, 82 

Fucus, 53, 91, 93 

Fucus vesciculosus, 93 

Fuligo septica, 177 

Fungi, 39, 69, 76, 110, 119, 146, 150, 172 

Fungi, bird's-nest, 155 

Fungi imperfecti, 140 

Fungilli, 39, 206 

Furcellariea, 46, 50 

Furcellarieae, 50 

Fusarium, 142 

Fusiformis, 29 

Fusobacterium, 29 

Fusulina, 188 

Fusulinida, 188 

Fusulinidae, 188 

Galaxaura, 47 
Galera tenera, 153 
Gallionella, 27 
Gallowaya, 148 
Gammarus, 233 
Gamocystis, 217 
Ganymedes, 216 
Ganymedida, 216 
Ganymedidae, 216 
Gasteromycetes, 152 
Gastrobionta, 6 
Gastrocarpeae, 50 
Geaster, 155 
Gelidiaceae, 49 
Gelidialea, 46, 49, 50 
Gelidiales, 49 
Gelidieae, 49 
Gelidium, 50, 51 
Geophonus, 186, 187 
Geoglossacea, 135 
Giardia, 163, 166 
Giardia enterica, 164, 166 
Giardia Lamblia, 166 
Gibberella, 142 
Gigantomonas, 167 
Gigartina mammilosa, 49 
Gigartinales, 47 
Gigartineae, 47 
Gigartininae, 47 
Glandulina, 187 
Glaucocystis, 33 
Glaucoma, 229 
Glaucoma pyriformis, 227 

Glenodinium, 94, 103 
Globigerina, 184, 188 
Globigerinida, 188 
Globigerinidea, 183, 187 
Globorotalia, 187 
Globorotaliidae, 187 
Gloeocapsa, 33 
Gloeochaete, 33 
Gloeochrysis, 62 
Gloeodiniaceae, 100 
Gloeodinium, 100 
Gloeosporium, 139, 140, 141 
Gloeotrichia, 36 
Gloiophycea, 31, 32, 33 
Gloiophyceae, 29, 33 
Glomerella, 126, 127, 139, 140 
Glomerella cingulata, 139 
Glugea, 222 
Glugeida, 222 
Glugeidae, 222 
Glugeidea, 222 
Glugeidees, 222 
Goat, 210 

Gomphonema, 72, 75 
Gomphonemaceae, 75 
Gomphonemea, 75 
Gomphonemeae, 75 
Gomphosphaeria, 33 
Gonapodiaceae, 112 
Gonapodiineae, 112 
Gonapodya, 112 
Goniaulax, 103 
Gonimophyllum, 52 
Goniodoma, 103 
Goniostomum, 109 
Goniotrichaceae, 43 
Goniotrichopsis, 43 
Goniotrichum, 43 
Gonococcus, 20 
Gonospora, 216 
Gooseberries, 146 
Goussia, 209, 210 
Goussia Schubergi, 207, 208, 237 
Gracilaria, 49 
Grains, 149 
Granuloreticulosa, 179 
Graphidiacea, 134 
Graphidiaceae, 134 
Graphidiales, 133 
Grasses, 149 

Green algae, see Algae, Green 
Gregarina, 206, 217 
Gregarina conica, 217 
Gregarina cuneata, 217 
Gregarina ovata, 217 
Gregarinae, 206, 216 
Gregarinarien, 217 
Gregarines 206, 209, 215, 219 
Gregarinida, 207, 217 
Gregarinidae, 217 
Gregarinidia, 207 
Gregarininea, 217 


The Classification of Lower Organisms 

Gregarinoidae, 217 
Gregarinoidea, 217 
Gregarinomorpha, 207 
Gromia, 179, 191 
Gromida, 191 
Guepinia, 150 
Guepinia apathularia, 145 
Gurleya, 222 
Guttulina, 203 
Guttulina sessilis, 204 
GuttuHnacea, 201, 203 
Guttulinaceae, 203 
GuttuHneae, 203 
Guttulineen, 203 
Guttulinidae, 203 
Guttulinopsis, 203 
Gymnamoebae, 201 
Gymnamoebida, 201 
Gymnascales, 130 
Gymnoascaceae, 130 
Gymnoascus, 131 
Gymnocraspedidae, 67 
Gymnodiniacea, 99, 100 
Gymnodiniaceae, 100 
Gymnodlniales, 99 
Gymnodinida, 100 
Gymnodinidae, 100 
Gymnodiniidae, 100 
Gymnodinina, 99 
Gymnodinioidae, 99 
Gymnodinium, 100 
Gymnodinium Lunula, 101, 104 
Gymnodinium striatum, 104 
Gymnogongrus, 49 
Gymnosporangium, 143, 148 
Gymnosporidae, 211 
Gymnosporidiida, 209, 211 
Gymnostomata, 229 
Gymnostomataceae, 229 
Gymnostomina, 229 
Gyrodinium, 101 
Gyromonas, 166 
Gyrophragmium, 152 
Gyrosigma, 75 

Haemamoeba, 213 
Hacmamoeba malariac, 213 
Haemamoeba vivax, 213 
Hacmogregarina, 2 1 1 
Haemogregarinida, 211, 212 
Haemogregarinidac, 211 
Haemogrcgarinina, 2 1 1 
Haemoproteidae, 212 
Haemoproteus, 213 
Haemoproteus Columbae, 212, 213 
Hacmosporidae, 211 
Hacmosporidia, 207, 211, 212 
Hacmosporidiida, 211 
Haliarchnion, 49 
Halicryptina, 198 
Haliomma, 195 

Haliomma capillaris, 196 
Haliommatina, 195 
Halkyardia, 187 
Halopappaceae, 60 
Halopappus, 60 
Halosphaeraceae, 66 
Halteria, 231 
Halteridiida, 212 
Halteridiidae, 212 
Halteridium, 212 
Halteriidae, 231 
Halterina, 231 
Hantkenina, 187 
Hantkeninidae, 187 
Hantschia, 75 
Haploactinomyxidae, 222 
Haplobacteriacei, 18 
Haplocyta, 215 
Haplodinium, 99 
Haplospora, 87 
Haplosporangium, 124 
Haplosporangium lignicola, 122 
Haplosporidia, 218 
Haplosporidies, 218 
Haplosporidiida, 218 
Haplosporidiidae, 218 
Haplosporidiidea, 209, 218 
Haplosporidium, 218 
Haplostichinae, 82 
Haplozoonidae, 102 
Hauerinina, 185 
Hedriocystis, 194 
Helicosorina, 185 
Heliodiscus, 195 
Heliodiscus Phacodiscus, 196 
Heliolithae, 58 
Helioflagellida, 189 
Helioflagellidae, 191 
Heliozoa, 63, 157, 189, 190, 205 
Heliozoariae, 189, 190 
Heliozoida, 189 
Helminthocladeae, 47 
Helminthosporium, 142 
Helotiacea, 135 
Helvellacea, 135 
Helvellales, 134 
Helvellineae, 134 
Hemiascales, 130 
Hemiasceae, 130 
Hemiasci, 129 
Hemiascineae, 130 
Hemibasidii, 145, 149 
Hcmicristcllaria, 187 
Hemicyclomorpha, 18 
Hcmidinium, 101 
Hemileia vastatrix, 148 
Hemisphaeriaceae, 134 
Hcmisphaeriales, 133 
Hemitrichia, 177 
Hemitrichia intorta, 176 
Hemophilus, 22 
Henneguya, 221 



Hepatozoon, 211 
Herpetomonas, 161, 162 
Heterocapsaceae, 65 
Heterocapsales, 63 
Heterocapsineae, 55, 63 
Heterocarpea, 41, 44, 52 
Heterocarpeae, 40, 44 
Heterochlorida, 63 
Heterochloridaceae, 66 
Heterochloridae, 66 
Heterochloridales, 63 
Heterochloridea, 63 
Heterochloridineae, 55, 63 
Heterochromonas, 59 
Heterococcales, 63 
Heterococcineae, 55, 63 
Heterodermaceae, 175 
Heterodermeae, 171 
Heterogeneratae, 82 
Heterohelicida, 188 
Heterohelicidae, 188 
Heterohelix, 188 
Heterokonta, 11, 55, 83 
Heterokontae, 53, 55, 63 
Heteromastigoda, 158 
Heteromonadina, 59 
Heteronema, 109 
Heteronemidae, 108 
Heterophryida, 191, 193 
Heterophryidae, 193 
Heterophrys, 193 
Heterosiphonales, 63 
Heterosiphoneae, 55, 63 
Heterostegina, 188 
Heterotricha, 228, 230 
Heterotrichaceae, 230 
Heterotrichales, 63 
Heterotrichida, 230 
Heterotrichina, 230 
Heterotrichineae, 55, 63 
Hexacontium, 195 
Hexaconus, 197 
Hexactinomyxon, 222 
Hexamastix, 167 
Hexamastix Termopsidis, 164 
Hexamita, 163, 166 
Hexamitidae, 166 
Hirmocystis, 217 
Hodotermitidae, 167 
Hoferellus, 221 
Holocyclomorpha, 18 
Holomastigotoides, 169 
Holomastigotoidida, 169 
Holomastigotoididae, 169 
Holophrya, 229 
Holophryidae, 229 
Holotricha, 228, 229 
Holotrichida, 229 
Homalogonata, 69 
Homo sapiens, 6 
Honey bees, 222 
Hoplonympha, 169 

Hoplonympha natator, 170 
Hoplonymphida, 169 
Hoplonymphidae, 169 
Hoplorhynchus, 218 
Hordeum vulgare, 6 
Hormogonales, 34 
Hormogoneae, 34 
Horse, 231 
Hyalobryon, 60 
Hyalodiscida, 201, 202 
Hyalodiscidae, 202 
Hyalodiscus, 202 
Hyaloklossia, 211 
Hyaloria, 149 
Hyalospora, 217 
Hydnacea, 151 
Hydnaceae, 151 
Hydnangiacea, 155 
Hydnangiaceae, 155 
Hydnei, 151 
Hydnum, 151 
Hydra, 203, 233 
Hydramoeba, 203 
Hydrocoleum, 35 
Hydrogenomonas, 24 
Hydruracea, 61, 62 
Hydruraceae, 62 
Hydruridae, 62 
Hydrurina, 62 
Hydrurus, 61 
Hydrurus foetidus, 56, 62 
Hyella, 36 

Hymenogastraceae, 155 
Hymenogastrales, 152 
Hymenogastrea, 155 
Hymenogastrei, 155 
Hymenogastrineae, 152 
Hymenomonadacea, 58, 60 
Hymenomonadaceae, 60 
Hymenomonadidae, 60 
Hymenornonas, 60 
Hymenomycetales, 150 
Hymenomycetes, 150 
Hymenomycetineae, 150 
Hymenothecii, 150 
Hymenostomata, 229 
Hyperammina, 183 
Hyperamminidae, 183 
Hypermastigida, 168 
Hypermastigina, 158, 166, 168 
Hyphochytriacea, 69 
Hyphochytriaceae, 69, 117 
Hyphochytrialea, 57, 61, 69, 70, 1 11 
Hyphochytriales, 69 
Hyphochytrium, 69, 117 
Hyphochytrium catenoides, 70 
Hyphomycetes, 121, 140, 141 
Hypnodiniaceae, 100 
Hypocreaceae, 137 
Hypocreales, 137, 138, 139, 142 
Hypodermia, 146, 147 
Hypodermii, 147 


The Classification of Lower Organisms 

Hypomyces, 142 

Hypomyces Solani var. Cucurbitae, 126, 

Hypotricha, 228, 232, 233 
Hypotrichaceae, 233 
Hypotrichida, 233 
Hypotrichina, 233 
Hysterangiacea, 155 
Hysterangiaceae, 155 
Hysteriacea, 134 
Hysteriaceae, 133, 134 
Hysteriales, 133, 141 
Hysteriineae, 133, 134 
Hysterophyta, 119 

Ichthyophthirius, 229 

Ichthyosporidium, 219 

Imperforida, 183 

Infusoires, 223 

Infusoires suceurs, 235 

Infusoria, 2, 37, 95, 118, 223, 228, 232, 

Inoperculata, 135 
Inophyta, 39, 119 
Insects, 69, 113, 117, 118, 124, 125, 155, 

159, 161, 165, 167, 216, 217, 220 
Invertebrates, 161, 210, 211, 215, 216 
lodamoeba, 203 
Irish moss, 49 
Irpex, 151 
Isoachlya, 79 
Isocarpeae, 69 
Isochrysidaceae, 59 
Isochrysidae, 59 
Isochrysidales, 57 
Isogeneratae, 82 
Isospora, 210 

Janczewskia, 52 
Jarrina, 210 
Joenia, 169 
Joeniidae, 169 
Joeniidea, 168 
Jocnina, 169 
Joenopsis, 169 
Jola, 147 
Junipers, 148 

Kalotermes, 169 
Kalotermitidae, 167 
Kalotcrmitinac, 166, 168 
Karyamocbina, 203 
Karyolysus, 2 1 1 
Kelps, 82, 83, 89, 90 
Keramosphaera, 185 
Keramosphacridac, 185 
Keramosphaerina, 185 
Kerona, 233 
Klebsiella (bacterium), 7, 22 

Klebsiella pneumoniae, 22 
Klebsiella (flagellate), 7 
Klebsiella alligata, 106 
Klossia, 211 
Klossiella, 211 
Kofoidia, 168, 169 
Kofoidiida, 169 
Kofoidiidae, 169 
Kurthia, 21 
Kurthiacea, 19, 21, 237 

Laboulbenia, 140 
Laboulbenia Guerinii, 140 
Laboulbenia Rougetii, 140 
Laboulbeniaceae, 140 
Laboulbenialea, 129, 140 
Laboulbeniales, 140 
Laboulbenieae, 140 
Laboulbeniineae, 140 
Laboulbeniomycetes, 140 
Labyrinthula, 203, 204 
Labyrinthula macrocystis, 203 
Labyrinthulida, 201, 203 
Labyrinthulidae, 203 
Lachnea scutellata, 127, 136 
Lachnobolus, 176 
Lacrymaria, 229 
Lactobacillaceae, 20 
Lactobacillus, 20 
Lactobacteriaceae, 20 
Lagena (oomycete), 82 
Lagena (rhizopod), 82, 184, 187 
Lagenaceae, 187 
Lagenidae, 186 
Lagenidea, 185 
Lagenidiacca, 81, 82 
Lagenidiaceae, 82 
Lagenidialea, 76, 81, 111, 118 
Lagenidiales, 81 
Lagenidium, 82 
Lagenina, 186 
Lagenocystis, 82, 237 
Lagenocystis radicicola, 82, 237 
Lagynida, 191 
Lagynion, 63 
Lagynis, 191 
Laminaria, 91 
Laminaria yczoensis, 92 
Laminariaccae, 89 
Laminariales, 89 
Laminariea, 85, 89 
Laminarieae, 89 
Lampoxanthium, 195 
Lampramocbac, 205 
Ivamprodernia, 1 75 
Lamprodermaceae, 1 75 
Lamprospora Iciocarpa, 136 
Lamprosporalcs, 171 
Lankestcria, 216 
Larcarida, 195 
Larcoidea, 195 



Latrostium, 69 

Laurencia, 52 

Leangium, 177 

Leathesia, 88 

Lecudina, 216 

Lecudinidae, 216 

Leeches, 161, 211 

Legerella, 211 

Leidyopsis, 169 

Leishmania, 162 

Leishmania brasiliensis, 162 

Leishmania Donovani, 162 

Leishmania tropica, 162 

Lemanea, 47 

Lemna, 69 

Lenticulina, 187 

Lenticulites, 187 

Lentospora, 221 

Lenzites, 151 

Leocarpus, 177 

Leocarpus fragilis, 176 

Lepidoderma, 177 

Lepidoderma Chailletii, 176 

Lepochromulina, 62 

Leptodiscida, 100, 102 

Leptodiscidae, 102 

Leptodiscus, 102 

Leptolegnia, 79 

Leptomitaceae, 79 

Leptomi tales, 77 

Leptomitea, 77, 79 

Leptomiteae, 79 

Leptomitus, 79 

Leptomonas, 162 

Leptospira, 29 

Leptospira icteroides, 29 

Leptospira icterohaemorrhagiae, 29 

Leptospironympha, 169 

Leptostromatacea, 141 

Leptostromataceae, 141 

Leptotheca, 221 

Leptothrix, 27 

Leptothrix ochracea, 27, 36 

Leptotrichacea, 27 

Leptotrichaceae, 20 

Leptotrichacei, 20, 27 

Leptotrichia, 20 

Leucocytozoidae, 212 

Leucocytozoon, 213 

Leuvenia, 66 

Liagora tetrasporifera, 47, 49 

Licea, 175 

Liceacea, 173, 175 

Liceaceae, 175 

Liceales, 171 

Liceidae, 175 

Lichenes, 119 

Lichens, 119, 120 

Ligniera, 179 

Lindbladia, 175 

Lionotus, 230 

Listeria, 21 

Lithocampe, 198 
Lithochytridina, 198 
Lithocircus, 198 
Lithocircus productus, 196 
Lithocolla, 193 
Lithocollidae, 193 
Lithocyclia, 195 
Lithocyclidina, 195 
Lithocystis, 216 
Litholophida, 197 
Litholophus, 197 
Lituola, 186 
Lituolidaceae, 186 
Lituolidae, 186 
Lituolidea, 185, 186 
Lituolina, 186 
Liverworts, 10 
Lizards, 211 
Lobosa, 201 
Loborhiza, 117 
Lobster, 211 
Loftusia, 186 
Loftusiidae, 186 
Loftusiina, 186 
Lophomonadidae, 169 
Lophomonadida, 168, 169 
Lophomonadina, 168 
Lophomonas, 168, 169 
Loxodes, 230 
Lychnaspis, 197 
Lycogala, 172, 175 
Lycogala epidendrum, 1 76 
Lycogalaceae, 171, 175 
Lycogactida, 174, 175 
Lycogalactidae, 175 
Lycogalales, 171 
Lycogalopsis, 155 
Lycogalopsis Solmsii, 145 
Lycoperdacea, 155 
Lycoperdaceae, 155 
Lycoperdales, 152 
Lycoperdineae, 152 
Lycoperdon, 155 
Lyngbya, 13, 35 
Lytothecii, 152 

Macrocystis pyrifera, 90, 91 

Macromastix, 58 

Macrotrichomonas, 167 

Macrotrichomonas pulchra, 164 

Maize, 6 

Mallomonadidae, 62 

Mallomonadinea, 61, 62 

Mallomonas, 61, 62 

Mallomonas roseola, 56 

Mammals, 166, 210 

Man, Mankind, Men, 6, 159, 165, 210, 

Margarita, 176 
Margaritaceae, 176 
Margaritida, 174, 176 


The Classification of Lower Organisms 

Margaritidae, 176 
Massospora, 124, 125 
Mastigamoeba, 158, 163 
Mastigamoeba aspera, 160 
Mastigamoebidae, 163 
Mastigella, 163 
Mastigophora, 6, 55, 94, 95 
Mastotermitidae, 167 
Matthewina, 186 
Mayorella, 202 
Mayorellida, 201, 202 
Mayorellidae, 202 
Medusetta, 200 
Medusettida, 200 
Megachytriaceae, 118 
Megachytrium, 118 
Melampsora, 143 
Melampsoracea, 148 
Melampsoraceae, 148 
Melanconiacea, 141 
Melanconiaceae, 141 
Melanconialea, 141 
Melanconiales, 141 
Melanophycea, 11, 55, 82 
Melanophyceae, 82 
Melanospermeae, 82 
Melitangium, 28 
Melosira, 72, 73, 74 
Melosiraceae, 74 
Melosireae, 74 
Meningococcus, 20 
Menoidium, 103, 107, 109 
Menoidium incurvum, 108 
Menospora, 218 
Menosporida, 218 
Menosporidae, 218 
Meridiea, 75 
Meridieae, 75 
Meridion, 75 
Meridionaceae, 75 
Merismopedia, 33 
Merocystis, 211 
Merogregarina, 215 
Merogregarinida, 215 
Merogregarinidae, 2 1 5 
Merolpidiaceae, 1 1 7 
Meroselenidium, 215 
Mesocaena, 63 
Mesogloia, 88 
Mesogloiacca, 88 
Metachaos, 202 
Metadevescovina, 167 
Metaphyta, 6 
Metasporeae, 1 1 7 
Metazoa, 6 
Metchnikovclla, 219 
Metchnikovellida, 219 
Mctchnikovellidae, 219 
Methanomonas, 24 
Micrococcacea, 19, 20 
Micrococcaceae, 20 
Micrococcus, 20 

Microcoleus, 35 
Microglena, 62 
Micromycopsis, 117 
Micropeltidacea, 134 
Micropeltidaceae, 134 
Microrhopalodina, 166 
Microsphaera, 132, 133 
Microsphaera alni, 133 
Microsporidia, 222 
Microsporidies, 222 
Microthyriacea, 134, 141 
Microthyriaceae, 134 
Microthytriales, 133 
Miescher's tubes, 206 
Mieschersche Schlauche, 206, 214 
Mikrogromia, 183 
Miliola, 182, 185, 201 
Milioles, 179 
Miliolida, 185 
Miliolidae, 185 
Miliolidea, 183, 185 
Miliolina, 185 
Mindeniella, 79 
Mischococcacea, 65, 66 
Mischococcaceae, 66 
Mischococcus, 66 
Mites, 211 
Mitraspora, 221 
Mitrati, 134 
Molds, 142 
Molds, water, 77 
Mollisiacea, 135 
Monaden, 189 
Monades, 59 
Monadidae, 59, 60 
Monadidea, 158 
Monadina, 57, 58, 59, 158 
Monadineae Tetraplasteae, 191 
Monadineae Zoosporeae, 191 
Monads, collared, 38 
Monas, 38, 54, 59, 60, 158 
Monas amyli, 189 
Monas Okenii, 31 
Monascus, 131 
Monera, 6, 12 
Moneres, 12, 189 
Monilia, 135, 140, 142 
Monilia sitophila, 139 
Moniliacea, 142 
Moniliaceae, 142 
Moniliales, 141 
Monkeys, 213 
Monoblepharella, 112 
Monoblepharella Taylori, 114 
Monoblcpharidacca, 112 
Monoblcpharidaceae, 112 
Monoblepharidalea, 111, 114 
Monoblepharidales, 1 1 1 
Monoblcpharideae, 110 
Monoblcpharidineae, 110, 111 
Monoblepharis, 111, 112 
Monocercomonadida, 167 



Monocercomonadidae, 167 
Monocercomonas, 167 
Monocercomonoides, 165 
Monocilia, 66 
Monociliaceae, 66 
Monocystid gregarines, 209 
Monocystida, 216 
Monocystidae, 216 
Monocystidea, 209, 215' 
Monocystiden, 216 
Monolpidiaceae, 118 
Monomychota, 17 
Monopylaria, 190, 196, 198 
Monopylea, 198 
Monopyleen, 198 
Monopylida, 198 
Monopylina, 198 
Monoschizae. 215 
Monosiga, 67, 68 
Monosomatia, 179, 183 
Monosporea, 210 
Monosporees, 209 
Monostomina, 191 
Morchella, 135 
Morchella conica, 136 
Mortierella, 124 
Mortierellacea, 123, 124 
Mortierellaceae, 124 
Mosquitoes, 162, 213 
Moss, Irish, 49 
Mosses, 10 

Mouse, Mice, 211,214 
Mrazekia, 222 
Mrazekiida, 222 
Mrazekiidae, 222 
Mucedinaceae, 142 
Mucedineae, 142 
Mucedines, 129, 130, 135 
Mucor, 121, 123 
Mucor Mucedo, 121, 123 
Mucoracea, 123 
Mucoraceae, 123 
Mucorales, 121 
Mucorina, 121, 128 
Mucorineae, 121 
Mucorini, 121 
Mucronina, 188 
Mushrooms, 145, 151 
Mussels, 211,218 
Mutinus, 155 
Mycetalia, 119 
Mycetoideum, Regnum, 119 
Mycetosporidium, 179 
Mycetozoa, 119, 157, 171, 176, 203 
Mycetozoen, 171, 172 
Mycetozoida, 171 
Mychota, 1,4,6,8, 10, 12 
Mycobacteriacea, 25 
Mycobacteriaceae, 25 
Mycobacterium, 25 
Mycobacterium leprae, 25 
Mycobacterium tuberculosis, 25 

Mycochytridinae, 113 
Mycoderma mesentericum, 24 
Mycophyceae, 77 
Mycophyta, 119 
Mycoporacea, 139 
Mycosphaerella, 139 
Mycosphaerella personata, 138 
Myrioblepharis, 112 
Myriogloiacea, 88 
Myrionema, 89 
Myrionematacea, 88 
Myriospora, 2 1 1 
Myxidiea, 221 
Myxidiees, 221 
Myxidiida, 221 
Myxidiidae, 221 
M>Tcidium, 221 
Myxobacter, 28 
Myxobacteria, 12, 14 
Myxobacteriacea, 28 
Myxobacteriaceae, 27, 28 
Myxobacter iales, 27 
Myxobactralea, 26, 27 
Myxobactrales, 27 
Myxobolea, 221 
Myxobolees, 221 
Myxobolida, 221 
Myxobolidae, 221 
Myxobolus, 221 
Myxoceratida, 221, 237 
Myxoceros, 221, 237 
Myxoceros Blennius, 220, 221, 237 
Myxoceros sphaerulosa, 221, 237 
Myxochloridae, 66 
Myxochrysidaceae, 63 
Myxochrysidae, 63 
Myxochrysis, 63 
Myxochytridinae, 113 
Myxococcacea, 28 
Myxococcaceae, 28 
Myxococcus, 28 
Myxococcus coralloides, 26 
Myxocystoda, 99 
Myxogastres, 171 

Myxomycetes, 10, 157, 171, 172, 178 
Myxomyceten, 172 
Myxomycidium flavum, 143 
Myxomycophyta, 171 
Myxophyceae, 17, 29, 30 
Myxophykea, 29 
Myxophyta, 171 
Myxoproteus, 221 
Myxoschizomycetae, 27 
Myxoschizomycetes, 18, 27 
Myxosoma, 221 
Myxosomatida, 221 
Myxosomatidae, 221 
Myxosporidia, 206, 219, 220 
Myxothallophyta, 171 
Myzocytium, 82 


The Classification of Lower Organisms 

Naegelliella, 62 
Naegelliellaceae, 62 
Naegelliellidae, 62 
Naegleria. 159 
Najadea, 60 
Nassellaria, 198 
Nassellida, 198 
Nassoidea, 198 
Nassula, 230 
Nassulidae, 230 
Nautilus, 182, 186, 187 
Navicula, 72, 73, 75 
Naviculaceae, 75 
Naviculales, 74 
Naviculea, 75 
Naviculeae, 75 
Neactinomvxon, 222 
Nebela, 205 
Nebelida, 205 
Nebelidae, 205 
Nectria, 141, 142 
Nectria cinnabarina, 139 
Nectrioidaceac, 141 
Nectrioideae, 141 
Neisseria gonorrhoeae, 20 
Neisseria intracellularis, 20 
Neiseria meningitidis, 20 
Neisseria Weichselbaumii, 20 
Neisseriacea, 19, 20 
Neisseriaceae, 20 
Neisseriacees, 20 
Nemalion, 47 
Nemalion multifidum, 49 
Nemalionales, 47 
Nemalioninae, 47 
Nemastomatales, 47 
Nematochrysidaceae, 60 
Nematochrysis, 61 
Nematocystida, 219 
Nematodes, 113, 118, 124 
Nematothecia, 141 
Nematothecii, 141 
Neogregarina, 215 
Neosporidia, 206, 207, 219 
Nephroselmidacea, 98 
Nephrosclmidaceae, 98 
Nephrosclmidae, 98 
Nephroselmis, 98 
Nereocystis, 89 

Nereocystis Luetkeana, 90, 91 
Neurospora, 139, 140 
Neurospora crassa, 127 
Neusinidae, 186 
Neusina, 186 
Nevskia, 27 
Nidularia, 155 
Nidulariaceae, 155 
Nidularialcs, 152 
Nidularica, 155 
Nidularici, 155 
Nidulariincae, 152 
Nina, 217,218 

Nitrobacter, 24 
Nitrobacter Winogradskyi, 24 
Nitrobacteriacea, 20, 24 
Nitrobacteriaceae, 24 
Nitromonas, 24 
Nitrosococcus, 24 
Nitrosococcus nitrosus, 24 
Nitrosomonas europaea, 24 
Nitrosomonas javanensis, 24 
Nitzschia, 75 
Nitzschiacea, 75 
Nitzschiaceae, 75 
Noctiluca, 95, 99, 102 
Noctiluca miliaris, 102 
Nectiluca scintillans, 102, 104 
Noctilucae, 94, 99 
Noctilucida, 100, 102 
Noctilucidae, 102 
Nodosalida, 186 
Nodosarella, 188 
Nodosaria, 184, 187 
Nodosarida, 186 
Nodosaridae, 186 
Nodosarina, 186, 188 
Nodosaroum, 186 
Nodosinella, 186 
Nodosinellida, 186 
Nodosinellidae, 186 
Nonion, 184, 187 
Nonionidea, 187 
Nonionideae, 187 
Nonionina, 187 
Nosema, 222 

Noscma bombycis, 206, 222 
Nosematidae, 222 
Nostoc, 35 
Nostocacea, 34, 35 
Nostocaceae, 35 
Nostochineae, 33 
Nowakowskiella, 118 
Nowakowskiellacea, 117, 118 
Nowakowskiellaceae, 118 
Nubecularina, 185 
Nucleophaga, 118 
Nuda, 201 
Nummulitaceae, 188 
Nummulites, 188 
Nummulitida, 188 
Nummulinidae, 188 
Nunmiulitina, 188 
Nummulitinidea, 183, 185, 188 

Oats, 148 

Ochromonadaceae, 59, 60 
Ochromonadalea, 54, 56, 57, 61, 64, 67, 

85, 165 
Ochromonadales, 57 
Ochromonadidae, 59 
Ochromonas, 58, 59, 60 
Ochromonas granulans, 54 
Octomitus, 166 



Octomyxa, 179 
Octospora, 222 
Oicomonadacea, 159, 161 
Oicomonadaceae, 161 
Oicomonadidae, 161 
Oidium, 142 
Oikomonas, 161 
Oligochaet worms, 222 
Oligonema, 177 
Oligosporea, 209, 210 
Oligotricha, 230 
Oligotrichaceae, 230 
Oligotrichida, 230 
Oligotrichina, 230 
Olpidiacea, 115, 118 
Olpidiaceae, 118 
Olpidopsidacea, 81 
Olpidiopsidaceae, 81 
Olpidiopsis, 81 
Olpidium, 113, 118 
Olpidium Allomycetos, 116 
Ommatida, 195 
Onygena, l31 
Oodinidae, 102 

Oomycetes, 11, 53, 55, 65, 76, 78, 
118, 119, 121, 125, 127, 177, 178, 
Oosporeae, 77 
Opalina, 225, 227, 229 
Opalinalea, 228, 237 
Opalinida, 228 
Opalinidae, 228, 229 
Opalinina, 229 
Opalininea, 228 
Opalinoea, 225, 229 
Operculata, 135 
Operculina, 188 
Ophiocytiaceae, 66 
Ophiocytium, 66 
Ophiocytium parvulum, 66 
Ophiotheca, 176 
Ophrydium, 233 
Ophryocystis, 215 
Ophryocystida, 2 1 5 
Ophryocystidae, 215 
Ophryodendrida, 236 
Ophryodendridae, 236 
Ophryodendrina, 236 
Ophryodendron, 236 
Ophryoglena, 229 
Ophryoglenidae, 229 
Ophryoscolecidae, 231 
Ophryoscolecids, 225 
Ophryoscolccina, 231 
Ophryoscolex, 231 
Ophthalmidium, 184, 185 
Opisthokonta, 39, 110, 121, 237 
Opistokonten, 111 
Orbitoides, 188 
Orbitoidida, 188 
Orbitoididae, 188 
Orbitolina, 186 
Orbitolinida, 186 

Orbitolinidae, 186 
Orbitolites, 185 
Orbulina, 188 
Orbulinida, 188 
Orcadella, 175 
Orcadellaceae, 175 
Orcadellidae, 175 
Orcheobius, 211 
Orobias, 188 
Ortholithinae, 58 
Orthopteran, 217 
Orthosporeae, 117 
Oscillaria malariae, 213 
Oscillatoria, 30, 35, 36 
Oscillatoria Princeps, 13 
Oscillatoria splendida, 32 
Oscillatoriacea, 34, 35 
Oscillatoriaceae, 35 
Owl, 162 

Ox, Oxen, 162, 231 
Oxymonadida, 165, 166 
Oxymonadidae, 166 
Oxymonadina, 163 
Oxymonas, 163, 166 

111, Oxyphysis, 103 

179 Oxyrrhis, 101_ 

Oxyrrhis marina, 101 
Oxytocum, 103 
Oxytricha, 233 
Oxytrichidae, 233 
Oxytrichina, 233 

Pacinia, 23 

Pacinia cholerae-asiaticae, 23 

Padina, 87 

Palatinella, 62 

Pantostomatales, 158 

Pantostomatida, 158 

Pantostomatineae, 158 

Paradinida, 98 

Pardinidae, 98 

Paradinium Pouchetii, 97, 98 

Paraisotricha, 231 

Parajoenia, 167 

Paramaecium, 223, 224, 225, 226, 227, 

Paramaecium Aurelia, 226, 227 
Paramaecium Bursaria, 226 
Paramaecium caudatum, 226 
Paramaecium multimicronucleatum, 226 
Parameciina, 229 
Paramoeba Eilhardi, 98 
Paramoebida, 98 
Paramoebidae, 98 
Paramoecidae, 229 
Parasitella, 123 
Parvobacteriaceae, 22 
Pasteurella avicida, 22 
Pasteurella pestis, 22 
Pasteurellacea, 19, 22, 23, 237 
Pasteuria, 26, 27 


The Classification of Lower Organisms 

Patellariacea, 135 

Patellina, 181, 182, 185 

Patouillardina, 149 

Patouillardina cinerea, 145 

Pavonina, 188 

Peach, 137 _ 

Pectobacterium, 23 

Pectobacterium carotovorum, 23 

Pedangia, 186 

Pedilomonas, 111 

Pedinella, 62, 63 

Pegidia, 188 

Pegidiida, 188 

Pegidiidae, 188 

Pelodictyon, 31 

Pelomyxa, 202 

Pelomyxa carolinensis, 200, 201 

Pelomyxa palustris, 202 

Peneroplidae, 185 

Peneroplidea, 185 

PeneropHdina, 185 

Peneroplis, 181, 184, 185 

Penicillium, 130, 131 

Penicillium notatum, 25, 131 

Pennatae, 74 

Penta trichomonas, 165 

Pentatrichomonas obliqua, 164, 167, 237 

Peranema, 108, 109 

Peranema trichophorum, 108 

Peranemaceae, 108 

Peranemina, 108 

Perforida, 186 

Periblasteae, 47 

Perichaena, 176 

Perichaenacea, 174, 176 

Perichaonaceae, 176 

Peridinaca, 102, 103 

Peridinea, 96 

Peridineae, 94, 96, 103 

Peridiniaceae, 103 

Peridiniales, 102 

Peridinidae, 103 

Peridiriina, 103 

Peridinioidae, 103 

Peridinium, 94, 103 

Peridinium cinctum, 104 

Perionella, 66 

Pcripylaria, 194 

Pcripylea, 194 

Peripyleen, 194 

Peripylida, 194 

Peripylina, 194 

Perisporia, 131 

Porisporiacea, 129, 131 

Perisporiaceae, 131 

Perisporialcs, 131 

Peritricha, 233 

Peritrichaceae, 233 

Peritrichida, 233 

Peritrichinae, 18 

Pcritromidae, 233 

Peritromina, 233 

Peritromus, 233 
Peronospora, 81 
Peronosporacea, 80, 81 
Peronosporaceae, 81 
Peronosporales, 80 
Peronosporina, 76, 80 
Peronosporinae, 80 
Peronosporineae, 80 
Peziza, 127, 135 
Peziza domiciliana, 127 
Pezizacea, 135 
Pezizales, 134 
Pezizineae, 134 
Pestallozia, 141 
Pfeiflferella mallei, 22 
Phacidiaceae, 133, 134 
Phacidiacei, 133 
Phacidialea, 129, 133, 135 
Phacidiales, 133 
Phacidiea, 134, 141 
Phacidieae, 134 
Phacidiineae, 133, 134 
Phacus, 94, 106, 107 
Phaenocystes, 219 
Phaenocystida, 219 
Phaeocapsa, 98 
Phaeocapsaceae, 98 
Phaeocapsales, 96 
Phaeococcus, 98 
Phaeoconchia, 198, 199 
Phaeocystia, 198 
Phaeocystina, 199 
Phaeocystis, 58 
Phaeocystis globosa, 54 
Phaeodaria, 199 
Phacodariae, 198 
Phaeodermatium, 63 
Phaeogromia, 198, 199 
Phaeophyceae, 53, 82, 95 
Phacophycophyta, 53 
Phaeophyta, 39, 53 
Phaeoplakaceae, 98 
Phaeoplax, 98 
Phaeosphaera, 59 
Phacosphaeria, 190, 196, 198, 199 
Phaeosporales, 86 
Phaeosporcae, 82, 86 
Phaeothamnion, 61 
Phaeothamnionacea, 58, 60 
Phaeothamnionaccae, 60 
Phaeozoosporea, 85, 86, 87 
Phaeozoosporeac, 86 
Phagomyxa, 179 
Phalanastrriaccae, 67 
Phalanastcriidae, 67 
Phalanasterium, 67 
Phalanasterium digitatum, 68 
Phallaccae, 155 
Phallales, 152 
Phallineac, 152 
Phalloidca, 155 
Phalloidei, 155 



Phallus, 155 
Phlebotomus, 21 
Phleospora, 139 
Phlyctidiacea, 115, 117 
Phlyctidiaceae, 117 
Phlyctidium, 117 
Phlyctorhiza, 117 
Phoma, 141 
Phomaceae, 141 
Phomales, 141 
Phomatacea, 141 
Phomataceae, 141 
Phomatalea, 141 
Phomatales, 141 
Phormidium, 32, 35 
Phraginidium, 147, 148 
Phragmidium violaceum, 147 
Phycochromaceae, 29 
Phycomyces, 123, 124 
Phycomyces nitens, 122 
Phycomyceten, 76 
Phycomycetes, 76 
Phycomycophyta, 76 
Phyllactinia, 132, 133 
Phyllactinia corylea, 127 
Phyliophora, 49 
Phyllosiphon, 67 
Phyllosiphonacea, 67 
Phyllosiphinaceae, 67 
Physaraceae, 171, 177 
Fhysaraks, 171, 174 
Physarea, 174, 177 
Physaridae, 177 

Physarum, 177 

Physarum notabile, 176 

Physarum polycephalum, 176 

Physematium, 189, 195 

Physoderma, 115, 117 

Physodermataceae, 117 

Physomonas, 59 

Phytodiniacea, 99, 100 

Phytodiniaceae, 100 

Phytodinidae, 100 

Phytodinium, 100 

Phytomastigophorea, 55 

Phytomonas (bacterium), 7, 23 

Phytomonas (flagellate), 7, 161 

Phytomonas Donovani, 160 

Phytomyxida, 111, 171, 177 

Phytomyxidae, 179 

Phytomyxinae, 177 

Phytomyxini, 177 

Phytophthora, 80 

Phytophthora infestans, 81 

Phytosarcodina, 171 

Phytozoidea, 94, 105 

Pigeon, 212 

Pileati, 150 

Pileocephalus, 218 

Pilobolus, 121, 124 

Pinaciophora, 193 

Pinacocystis, 193 

Pines, 148 

Pinnularia, 72, 75 
Pipetta, 195 
Piptocephalidacea, 123, 124 

Piptocephalidaceae, 124 
Piptocephalis, 123, 124 

Piroplasma, 214 
Pisces, 1 

Plagiotomidae, 230 

Plagiotomina, 230 

Planopulvinulina, 187 

Planorbulina, 187 

Planorbulinidae, 187 

Plant kingdom, Plantae, Plants, 1,2, 4, 6, 
8, 10, 24, 38, 61, 67, 95, 113, 117, 118, 
130, 137, 148, 151, 161, 177, 179, 202 

Plasmodida, 213 

Plasmodidae, 213 

Plasmodiida, 211 

Plasmodiophora, 179 

Plasmodiophora Brassicae, 178 

Plasmodiophoraceae, 1 79 

Plasmodiophorales, 177 

Plasmodiophorea, 179 

Plasmodiophoreae, 179 

Plasmodiophoreen, 179 

Plasmodiophorina, 177 

Plasmodium, 212, 213 

Plasmodium falciparum, 214 

Plasmodium malariae, 213 

Plasmodium vivax, 213 

Plasmodroma, 157 

Plasmopara viticola, 81 

Platychrysis, 58 

Platygloea, 147 

Platynoblasteae, 51 

Platysporea, 221 

Plectascales, 130 

Plectascineae, 130 

Plectellaria, 198 

Plectida, 198 

Plectobasidiales, 152 

Plectobasidiineae, 152 

Plectofrondicularia, 188 

Plectoidea, 198 

Plectonema, 36 

Plectonida, 198 

Pleurage curvicolla, 128 

Pleurocapsa, 36 

Pleurocapsacea, 35, 36 

Pleurocapsaceae, 36 

Pleuromonas (dinoflagellate), 99 

Pleuromonas (zoomastigote), 159 

Pleuronemidae, 230 

Pleurosigma, 75 

Pleurostomella, 188 

Pleurostomellida, 188 

Pleurostomellidae, 188 

Pleurotricha, 233 

Pleurotrichidae, 233 

Pleurotus, 152 

Pleurotus ostreatus, 152 


The Classification of Lower Organisms 

Plistophora, 222 

Plistophoridae, 222 

Plocapsilina, 186 

Plocapsilinidae, 186 

Plowrightia morbosa, 140 

Pneumobacilliis, 22 

Podangium, 28 

Podaxacea, 152 

Podaxaceae, 152 

Podaxon, 152 

Podocyathus, 236 

Podophrya, 235 

Podophyridae, 235 

Podophryina, 235 

Podosphaera, 132, 133 

Polyangiaceae, 28 

Polyangidae, 27 

Polyangium, 28 

Polychaos, 202 

Polychytrium, 117 

Polycystidea, 209, 216 

Polycystina (of Ehrenberg), 189, 198 

Polycystina (of Delage and Herouard), 

Polydinida, 101 
Polygastrica, 223 
Polykrikida, 100, 101 
Polykrikos, 101 
Polymastigida, 158, 163, 164 
Polymastigidae, 165 
Polymastigina, 158, 163, 165 
Polymastix, 163, 165 
Polymastix melolonthae, 164 
Polymorphina, 187 
Polymorphinida, 187 
Polymorphinidae, 187 
Polymorphinina, 187 
Polymyxa, 178, 179 
Polyphagaceae, 117 
Polyphagus, 111, 117 
Polyphagus Euglcnae, 116, 117 
Polyporacea, 151 
Polyporaceae, 151 
Polyporales, 150 
Polyporei, 151 
Polyporus, 151 
Polysiphonia nigrescens, 49 
Polysiphonia violacea, 45, 46 
Polysiphonieae, 51 
Polysomatia, 179, 185 
Polysphondylium, 203 
Polysphondylium violaceum, 204 
Polysporea, 209,211 
Polystichinae, 82 
Polystictus, 151 
Polystomclla, 186, 187 
Polystomella crispa, 181 
Polystomellina, 187 
Polythalamia, 179, 185 
Polytoma, 61 
Pontifex, 202 
Pontisma, 81 

Pontosphaera, 60 
Pontosphaeraceae, 60 
Porospora, 218 
Porosporida, 218 
Porosporidae, 218 
Porphyra, 43 
Porphyra laciniata, 42 
Porphyra tenera, 42, 43 
Porphyra umbilicaris, 42, 43 
Porphyraceae, 43 
Porphyrea, 41, 43 
Porphyreae, 43 
Porphyridiacea, 41 
Porphyridiaceae, 41 
Porphyridiales, 4l 
Porphyridium, 3, 40 
Porphyridium cruentum, 41 
Postelsia palmaeformis, 90, 91 
Poteriochromonas, 60 
Poteriodendron, 67 
Poteriodendron petiolatum, 68 
Pouchetia, 101 
Pouchetiida, 100, 101 
Pouchetiidae, 101 
Prasiola, 3, 40, 44 
Prasiolaceae, 44 
Primalia, 37 

Primigenium, Regnum, 37 
Proboscoidella, 166 
Progastreades, 94, 95 
Pronoctiluca, 101 
Prorocentraceae, 99 
Prorocentrales, 99 
Proroccntridae, 99 
Proroccntrina, 99 
Prorocentrinea, 98 
Prorocentrinen, 99 
Prorocentrum, 99 
Prorodon, 229 
Protamoeba, 189 
Proteomyxa, 189, 190 
Proteomyxiae, 189, 190 
Proteomyxida, 189 
Proteromonadidae, 159 
Protcromonadina, 158 
Proteromonas, 159 
Proterospongia Haeckcli, 68 
Proteus diffluens, 201 
Proteus vulgaris, 22 
Protista, 4, 6, 37, 189 
Protistcs trichocystiferes, 94, 95 
Protoascineac, 130 
Protobasidiomycetes, 145, 146, 150 
Protobionta, 6, 37 
Protochrysis, 98 
Protociliata, 228 
Protoctista, 1, 4,6,8, 10,37 
Protodennieae, 171 
Protodinifcr, 101 
Protodiniferida, 100, 101 
Protodiniferidae, 101 
Protodiscineae, 137 



Protodontia Uda, 145 

Protoflorideae, 41 

Protogenes, 189 

Protomastigales, 158 

Protomastigida, 158 

Protomastigina, 158 

Protomastigineae, 158 

Protomonas, 189, 191 

Protomonadina, 158 

Protomyces, 130 

Protoopalina, 229 

Protoopalinidae, 229 

Protophyta, 6, 12, 18 

Protoplasta, 39, 111, 157 

Protoplasta filosa, l90 

Protopsis, 101 

Protozoa, 6, 12, 29, 37, 39, 223 

Prowazekia, 159 

Prunoidea, 195 

Prunophracta, 197 

Prymnesiidae, 58 

Prymneslum, 58 

Pseudomonas, 23 

Pseudomonas aeruginosa, 23 

Pseudospora, 159, 189, 191 

Pseudosporea, 19l 

Pseudosporeae, 191 

Pseudosporeen, 191 

Pseudosporidae, 191 

Pseudotetraedron, 66 

Pseudotetraedron neglectum, 64 

Psorosperms, 206 

Psychodiere, Regne, 37 

Psychodies, 37 

Pteridomonas, 193 

Pterocephalus, 218 

Pterospora, 216 

Ptychodiscida, 103 

Puccinia, 143, 147, 148 

Puccinia graminis, 147, 148 

Puccinia Malvacearum, 148 

Pucciniaceae, 148 

Pucciniales, 147 

Puffballs, 155, 172 

Punctariales, 89 

Pycnospermeae, 82, 89 

Pylaiella, 86 

Pyrenomycetales, 138 

Pyrenomycetes, 137 

Pyrenomycetineae, 137 

Pyrgo, 185 

Pyrocystis, 100 

Pyronema, 127, 134, 135,_ 137 

Pyronema confluens var. igneum, 127 

Pyronemacea, 135 

Pyrrhophycophyta, 94 

Pyrrhophyta, 39, 94, 182 

Pyrsonympha, 166 

Pyrsonymphina, 163 

Pythiacea, 80 

Pythiaceae, 80 

Quadrula, 205 

Rabbit, 210 
Raciborskya, 100 
Radaisia, 36 
Radioflagellata, 190 
Radiolaria, 189, 190, 194, 196 
Radiolariae, 189 
Radiolarida, 189 
Ralfsia, 87, 89 
Ralfsiacea, 88 
Ramularia, 139 
Ramulinina, 187 
Raphidophrys, 193 
Raphidozoum, 195 
Rat, 160 
Ravenelia, 148 
Red algae, see Algae, Red 
Regne Psychodiere, 37 
Regnum Mycetoideum, 119 
Regnum Primigenium, 37 
Reophacida, 186 
Reophacidae, 186 
Reophax, 186 
Reptiles, 212, 220 
Reticularia, 175, 179 
Reticulariacea, 174, 175 
Reticulariaceae, 175 
Reticularieae, 171 
Reticulitermes, 171 
Reticulosa, 179 
Retortomonadidae, 165 
Retortomonadina, 163 
Retortomonas, 163, 165 
Rhabdogeniae, 207 
Rhabdosphaera, 60 
Rhipidiacea, 77, 79 
Rhipidiaceae, 79 
Rhipidium, 79 
Rhizammina, 183 
Rhizamminidae, 183 
Rhizaster, 63 
Rhizidiacea, 115, 117 
Rhizidiaceae, 117 
Rhizidiomyces, 69 
Rhizidiomyces apophysatus, 70 
Rhizidiomycetaceae, 69 
Rhizidium, 113, 117 
Rhizinacea, 135 
Rhizo-Flagellata, 158 
Rhizobiacea, 19, 22, 23 
Rhizobiaceae, 22 
Rhizobium, 23 

Rhizobium Leguminosarum, 23 
Rhizochloridaceae, 66 
Rhizochloridae, 66 
Rliizochloridales, 63 
Rhizochloridea, 63 
Rhizochloridineae, 55, 63 
Rhizochloris, 66 
Rhizochrysidaceae, 63 


The Classification of Lower Organisms 

Rhizochrysidae, 63 

Rhizochrysidina, 61 

Rhizochrysidinae, 61 

Rhizochrysidineae, 55 

Rhizochrysis, 61, 63 

Rhizochrysis Scherffeli, 56 

Rhizocryptineae, 95 

Rhizoctonia, 142 

Rhizodiniales, 99, 101 

Rhizodininae, 95, 99 

Rhizoflagellata, 157, 158, 160, 178, 192 

Rhizomastigaceae, 163 

Rhizomastigida, 158 

Rhizomastigina, 158, 163 

Rhizomastix, 163 

Rhizopoda, 6, 63, 157, 172, 179, 184, 200, 

Rhizopoda radiaria, 189, 194 
Rhizopods, 179 
Rhizopodes, 179 
Rhizopogonacea, 155 
Rhizopogonaceae, 155 
Rhizopus, 121 

Rhizopus nigricans, 122, 124 
Rhizosolenia, 74 
Rhizosoleniacea, 74 
Rhizosoleniaceae, 74 
Rhodobacillacea, 31, 237 
Rhodobacillus, 31 
Rhodobacteria, 30, 31 
Rhodobacteriaceae, 31 
Rhodochaetacea, 41, 43 
Rhodochaetaceae, 43 
Rhodochaete, 43 
Rhodochorton, 47 
Rhodomelaceae, 51 
Rhodomeleae, 51 
Rhodomonas, 98 
Rhodomonas baltica, 97 
Rhodophyceac, 6, 40 
Rhodophycophyta, 40 
Rhodophyllis, 49 
Rhodophyta, 39, 40, 44 
Rhodopseudomonas, 31 
Rhodospermeae, 40 
Rhodospirillum, 31 
Rhodymcniacea, 51 
Rhodymeniaceae, 51 
Rhodymeniales, 51 
Rhodymcnieae, 51 
Rhodymeninae, 51 
Rhoicosphenia, 76 
Rhoicosphenia curvata, 72 
Rhopalodia, 75 
Rhynchocystida, 216 
Rhynchocystidae, 216 
Rhynchocystis, 216 
Rhynchomonas, 159 
Rickettsia Mclophagi, 21 
Rickettsia Prowazekii, 21 
Rickettsia Rickettsii, 21 
Rickettsiacea, 19, 20, 118 

Rickettsiaceae, 20 
Rivularia, 36 
Rivulariacea, 34, 36 
Rivulariaceae, 36 
Roach, 166, 168,170 
Rodents, 211 
Roesia, 69 
Rosaceae, 148 
Rotalia, 184, 187 
Rotaliaceae, 187 
Rotalida, 187 
Rotalidae, 187 
Rotalidea, 187 
Rotalina, 187 
Rotifers, 113, 118, 219 
Rozella, 118 
Rugipes, 202 
Rupertia, 187 
Rupertiidae, 187 
Russula, 143 
Russula emetica, 145 
Rusts, 145, 147 
Rye, 148 

Saccamminidae, 183 
Saccharomyces cerevisiae, 130 
Saccharomycetacea, 130 
Saccharomycetaceae, 130 
Saccharomycetes, 130 
Saccinobaculus, 163, 166 
Sagosphaerida, 199 
Sagrina, 188 
Salmonella, 22 
Salpingoeca, 67 
Salpingoeca ampullacea, 68 
Salpingoeca Clarkii, 68 
Salpingoecidae, 67 
Sappinia, 203 
Sappinia diploidea, 203 
Sappinia pedata, 204 
Sappiniaceae, 203 
Sappiniidae, 203 
Saprolegnia, 76, 79 
Saprolegnia ferax, 78 
Saprolegnia mixta, 78 
Saprolegniaceae, 77 
Saprolegniales, 77 
Saprolegniea, 77 
Saprolegnicae, 77 
Saprolcgniineae, 77 
Saprolegnina, 77 
Saprolegninae, 77 
Sapromyces, 79 
Saprospira, 29 
Sarcina, 20 
Sarcocystida, 214 
Sarcocystidae, 214 
Sarcocystidca, 214 
Sarcocystis, 214 
Sarcocystis Miescheriana, 214 
Sarcocystis Muris, 214 



Sarcodina, 6, 172, 200 

Sarcosporidia, 207, 214 

Sargassaceae, 91 

Sargassea, 92 

Sargasseae, 92 

Sargassum, 93 

Sargassum Horneri, 93 

Saricodina, 63, 157,200 

Schaudinella, 216 

Schaudinellida, 216 

SchaudinelHdae, 216 

Schinzia Leguminosarum, 23 

Schizocystida, 215 

Schizocystidae, 215 

Schizocystinea, 215 

Schizocystis, 215 

Schizodinium, 102 

Schizogoniacea, 41, 44 

Schizogoniaceae, 44 

Schizogonium, 44 

Schizogregarinaria, 215 

Schizogregarinida. 209, 215 

Schizomycetae, 17, 18 

Schizomycetes, 18, 206 

Schizomycophyta, 17 

Schizophyta, 12, 18 

Schizophytae, 12 

Schizosporea, 18 

Schlauche, Mieschersche, 206, 214 

Sciadiaceae, 66 

Sciadophora, 218 

Sclerocarpa, 129, 133, 135, 137, 145 

Sclerocarpi, 137 

Scleroderma, 143 

Sclerodermataceae, 155 

Sclerodermatales, 152 

Sclerodermea, 155 

Sclerodermei, 155 

Sclerotinia, 140 

Sclerotinia cinerea, 135, 136 

Scytomonas pusilla, 108 

Scytonema, 36 

Scytonematacea, 34, 35 

Scytonemataceae, 35 

Sebacina, 149 

Sebacina sublilacina, 145 

Sebdenia, 49 

Selenidium, 215 

Seleniida, 215 

Seleniidae, 215 

Selenococcidiida, 211 

Selenococcidiidae, 211 

Sclenococcidinea, 210 

Selenococcidium intermedium, 211 

Sennia, 97, 98 

Sepedonei, 141 

Septata, 217 

Septobasidium, 147 

Septoria, 139, 141 

Sheep, 210, 214 

Shigella, 22 

Shigella dysenteriae, 22 

Serratia, 22 

Siderocapsa, 27 

Sideromonas, 27 

Siedleckia, 215 

Silicina, 185 

Silicoflagellata, 55, 56, 57, 61, 62, 64, 67, 

Silicoflagellatae, 55, 62 
Silicoflagellidae, 62 
Silicoflagellina, 61 
Silkworms, 206, 222 
Sinuolinea, 221 
Siphonaria, 117 
Siphonogenerina, 188 
Siphonomycetae, 77 
Siphonophyceae, 55 
Siphonotestales, 62 
Sirolpidiacea, 81 
Sirolpidiaceae, 81 
Sirolpidium, 81 
Sirosiphon, 36 
Sirosiphonacea, 34, 36 
Sirosiphonaceae, 36 
Slavina, 222 
Smuts, 145, 149 
Snails, 161,211 
Snakes, 210 
Snyderella, 168 
Snyderella Tabogae, 164 
Solenodinium, l99 
Sorangiacea, 28 
Sorangiaceae, 28 
Sorangium, 28 
Soranthera, 89 
Sorites, 185 
Soritidae, 185 
Soritina, 185 
Sorodiscus, 179 
Sorophoreen, 203 
Sorosphaera, 179 
Sphacelaria, 86 
Sphacelarialea, 85, 86 
Sphacelariales, 86 
Sphacelariea, 86 
Sphacelarieae, 86 
Sphaeractinomyxon, 222 
Sphaerastrum, 193 
Sphaerellaria, 194 
Sphaeria, 138, 141 
Sphaeria Scirpi, 128 
Sphaeriaceae, 137 
Sphaeriales, 137, 138, 139, 141 
Sphaerida, 195 
Sphaeridea, 194 
Sphaerioidaceae, 141 
Sphaerioideae, 141 
Sphaerita, 118 
Sphaerobolacea, 155 
Sphaerobolaceae, 155 
Sphaerobolus, 155 
Sphaerocapsa, 197 
Sphaerocapsida, 197 


The Classification of Lower Organisms 

Sphaerocladia, 112, 113 

Sphaerococcales, 47 

Sphaerococcoidea, 46, 47, 50 

Sphaerococcoideae, 47 

Sphaeroeca, 67 

Sphaeroidea, 195 

Sphaeroidina (genus of Rhizopoda), 187 

Sphaeroidina (family of Radiolaria), 195 

Sphaeromyxa, 221 

Sphaerophracta, 197 

Sphaerophrya, 235 

Sphaeropsidales, 141 

Sphaeropsideae, 141 

Sphaerospora, 221 

Sphaerosporida, 221 

Sphaerosporidae, 221 

Sphaerosporea, 221 

Sphaerotheca, 127, 133 

Sphaerotheca pannosa, 133 

Sphaerotilacea, 33 

Sphaerotilaceae, 33 

Sphaerotilalea, 30, 33, 237 

Sphaerotilus, 30 

Sphaerotilus natans, 33 

Sphaerozoen, 194 

Sphaerozoida, 195 

Sphaerozoum, 189, 195 

Spirillacea, 19, 23 

Spirillaceae, 23 

SpirilHna, 181, 182, 185 

Spirillinidea, 185 

Spirillinina, 185 

Spirillum, 24 

Spirochaeta, 29 

Spirochaeta cytophaga, 26, 27 

Spirochaeta plicatilis, 28, 29 

Spirochaetacea, 29 

Spirochaetaceae, 29 

Spirochaetae, 27 

Spirochactalea, 28 

Spirochaetales, 28 

Spirochaets, 12, 14, 166, 167 

Spirochona, 233, 235 

Spirochonidae, 231 

Spirochonina, 230, 231 

Spirocystida, 215 

Spirocystidae, 215 

Spirocystidces, 215 

Spirocystis, 215 

Spirodinium, 231 

Spirodiscus, 66 

Spirodiscus fulvus, 64, 66 

Spirogyrales, 121 

Spirolina, 185 

Spironema, 222 

Spirophyllum, 27 

Spirostomum, 230 

Spirotricha, 230 

Spirotrichida, 230 

Spirotrichonympha, 168, 169 

Spirotrichonymphidae, 169 

Spirotrichonymyjhina, 168 

Spirulina, 35 
Sponges, 37, 67 
Spongocarpeae, 50 
Spongospora, 179 
Spongurida, 195 
Spongurus, 195 
Sporobolomyces, 145 
Sporochnales, 87 
Sporochnea, 88 
Sporochnoidea, 85, 87, 89 
Sporochnoideae, 87 
Sporochnus, 93 
Sporodinia, 124 
Sporochytriaceae, 117 
Sporomyxa, 179 
Sporozoa, 111, 206, 207, 219 
Sporozoans, 21, 162 
Sporozoaires, 207 
Sporozoaria, 206, 207 
Spumaria, 177 
Spumariaceae, 177 
Spumellaria, 194, 195 
Spyrida, 198 
Spyridieae, 51 
Spyridina, 198 
Spyroidea, 198 
Squamarieae, 50 
Squids, 210 
Staphylococcus, 20 
Staurocyclia, 195 
Staurojoenina, 169 
Staurojoenina assimilis, 170 
Staurojoeninida, 169 
Staurojoeninidae, 169 
Stelangium, 28 
Stemonitaceae, 171, 175 
Stemonitales, 171, 174 
Stemonitea, 174, 175 
Stemonitidae, 175 
Stemonitis, 175 
Stemonitis splendens, 176 
Stenophora, 217 
Stenophorida, 21 7 
Stenophoridae, 217 
Stentor, 227, 230 
Stentor coeruleus, 
Stentoridae, 230 
Stcntorina, 230 
Stephanida, 198 
Stephida, 198 
Stephoidea, 198 
Stereotestales, 62 
Stereum, 151 
Stictaceac, 134 
Stictea, 134 
Sticteac, 134 
Stictidaceac, 134 
Stictideae, 133 
Stigonema, 36 
Stigoncmataceae, 36 
Stiibaceae, 142 





Stilbeae, 142 
Stilbellacea, 142 
Stilbellaceae, 142 
Stilbosporei, 141 
Stilbum, 142 
Stilophora, 88 
Stilotricha, 233 
Stipitochloridae, 66 
Stipitococcacea, 65, 66 
Stipitococcaceae, 66 
Stipitococcus, 66 
Stokesiella, 60 
Stomaticae, 74 
Stomatoda, 223, 228, 233 
Stomatophora, 216 
Stomatophorida, 216 
Stomatophoridae, 216 
Streblomastigida, 165, 166 
Streblomastigidae, 166 
Streblomastix, 163, 168 
Streblomastix Strix, 164, 166 
Streblonema, 86 
Streptococcus, 20 
Streptomyces, 25 
Streptomycetaceae, 25 
Streptothrix, 25 
Striatae, 74 
Stylobryon, 60 
Stylocephalida, 218 
Stylocephalidae, 218 
Stylocephalus, 218 
Stylochrysalis, 59 
Stylocometes, 236 
Stylodinium, 100 
Stylonychia, 227, 232, 233 
Stylopage, 124 
Stylopyxis, 60 
Stylorhynchidae, 218 
Stypocaulon, 83, 84, 86 
Suctorea, 235 
Suctoria, 235 
Surirella, 71, 73, 75 
Surirella saxonica, 72, 73 
Surirellaceae, 75 
Surirellea, 75 
Surirelleae, 75 
Swine, 210, 214 
Symbelaria, 194 
Symploca Muscorum, 13 
Synactinomyxida, 222 
Synactinomyxidae, 222 
Synactinomyxon, 222 
Synchytriacea, 115, 117 
Synchytriaceae, 117 
Synchtrium, 117 
Syncephalastrum, 124 
Syncephalastrum racemosum, 122 
Syncephalis, 123, 124 
Syncephalis nodosa, 122 
Syncephalis pycnosperma, 122 
Syncollaria, 194 
Syncrypta, 59 

Syncryptaceae, 59 
Syncryptida, 58, 59 
Syncryptidae, 59 
Syncystida, 216 
Syncystidae, 216 
Syncystis, 216 
Syndinidae, 102 
Synedra, 72, 75 
Syntamiidae, 86 
Synura, 55, 59 
Synura Uvella, 54 
Synuraceae, 59 
Syracosphaera, 60 
Syracosphaera Quadricornu, 56 
Syracosphaeraceae, 60 
Syracosphaeridae, 60 
Syracosphaerinae, 57, 60 

Tabellaria, 75 

Tabellariaceae, 75 

Tabellariea, 75 

Tabellarieae, 75 

Taphrina, 127, 137 

Taphrina aurea, 137 

Taphrina deformans, 127, 136, 137 

Teliosporeae, 142 

Telomyxa, 222 

Telomyxa glugeiformis, 222 

Telomyxlda, 222 

Telomyxidae, 222 

Telosporidea, 207 

Telosporidia, 207 

Tentaculifera, 224, 228, 235 

Teratonympha, 171 

Teratonympha mirabilis, 170 

Teratonymphida, 169 

Teratonymphidae, 169 

Termites, 166, 167, 168, 169 

Termitidae, 168 

Termopsis, 166, 168 

Testacea, 205 

Testacida, 205 

Testaceolobosa, 205 

Tetractinomyxida, 222 

Tetractinomyxidae, 222 

Tetractinomyxon, 222 

Tetradinium, 100 

Tetradinium javanicum, 104 

Tetrahymena, 229 

Tetrahymena Geleii, 227 

Tetramitaceae, 165 

Tetramitida, 165 

Tetramitidae, 165 

Tetramitina, 165 

Tetramitus, 165 

Tetramyxa, 179 

Tetrasporeae, 82, 86 

Tetrasporees, 209 

Tetrataxis, 186 

Textularia, 182, 186 

Textulariaceae, 186 


The Classification of Lower Organisms 

Textularidae, 186 
Textularidea, 185 
Textularlna, 186 
Textulinida, 186 
Thalamophora, 179 
Thalassicolla, 189, 194, 195 
Thalassicollen, 194, 195 
Thalassicollida, 195, 199 
Thallochrysidacea, 62, 63 
Thallochrysidaceae, 63 
Thallochrysis, 63 
Thamnidium, 124 
Thaumatomastix, 109 
Thaumatomonadidae, 109 
Thaumatonema, 109 
Thaumatonemidae, 109 
Thecamoeba, 202 
Thecamoebae, 205 
Thecamoebida, 201, 202 
Thecamoebidae, 202 
Theileria, 214 
Theileridae, 214 
Thelephora, 151 
Thelephoracea, 151 
Thelephoraceae, 151 
Thelephorei, 151 
Thelohania, 222 
Theoconus, 198 
Thiere, 172 
Thiobacillus, 24 
Thiobacteria, 30, 31, 35 
Thiorhodaceae, 31 
Thioploca, 35 
Thiospira, 24, 31 
Thiospirillum, 31 
Thiothrix, 35 
Thoracosphaeraceae, 60 
Thoracosphaeridae, 60 
Thorea, 47 

Thraustochytriacea, 81, 82 
Thraustochytriaceae, 82 
Thraustochytrium proliferum, 82 
Thraustotheca, 79 
Ticks, 161, 206 
Tilletia, 149 
Tilletia Tritici, 145 
Tilletiacea, 149 
Tilletiaceae, 149 
Tilopteridales, 86 
Tilopteridca, 87 
Tilopterideae, 87 
Tilopteris, 87 
Timothy, 148 
Tinoporidea, 187 
Tinoporus, 187 
Tintinnidac, 231 
Tintinnids, 224 
Tintinnina, 231 
Tintinnodea, 231 
Tintinnoinea, 231 
Tipulocystis, 215 
Toads, 125 

Toadstools, 151 
Tokophrya, 235 
Tokophrya Lemnarum, 234 
Tolypothrix, 35, 36 
Torula, 130 
Torulopsis, 130 
Toxonema, 222 
Tracheliidae, 230 
Trachelina, 230 
Trachelius, 230 
Trachelomonas, 94, 106, 107 
Transchelia, 143 
Tremella, 149 
Tremella Auricula, 146 
Tremellacea, 149 
Tremellaceae, 149 
Tremellales, 149 
Tremellina, 146, 149, 150 
Tremellineae, 146, 149 
Tremellinei, 149 
Tremellini, 149 
Tremellodendron, 149 
Trepomonadida, 165, 166 
Trepomonadidae, 166 
Trepomonas, 166 
Treponema, 29 

Treponema macrodentium, 29 
Treponema microdentium, 29 
Treponema pallidum, 28, 29 
Treponema pertenue, 29 
Treponematacea, 29 
Treponemataceae, 29 
Tretomphalus, 180 
Triactinomyxon, 222 
Triactinomyxidae, 222 
Tribonema, 65, 66, 73, 95 
Tribonema bombycina, 64 
Tribonematacea, 65, 66 
Tribonemataceae, 66 
Triceratium, 74 
Tricercomitus, 167 
Tricercomitus Termopsidis, 164 
Trichamoeba, 202 
Trichia, 176, 177 
Trichiacea, 174, 176 
Trichiaceae, 171, 176 
Trichiales, 171, 174 
Trichiidae, 177 
Trichina, 177 
Trichinaceae, 1 76 
Trichoblasteae, 51 
Trichocystiferes, Protistes, 94, 95 
Trichodina, 235 
Trichodinidae, 235 
Trichomitus, 166 
Trichomonadida, 166, 167 
Trichomonadidae, 166, 167 
Trichomonadina, 158, 164, 166 
Trichomonads, 165 
Trichomonas, 166, 167 
Trichomonas hominis, 165 
Trichomonas tenax, 164, 167 



Trichomonas Termopsidis, 168 
Trichomonas vaginalis, 167 
Trichonympha, 168, 169, 170 
Trichonympha Campanula, 168, 170 
Trichonympha sphaerica, l68 
Trichonymphida, 169 
Trichonymphidae, 169 
Trichonymphidea, 168 
Trichonymphina, 168 
Trichophyton, 142 
Trichospermi, 152, 171 
Trichostomata, 229 
Tridictyopus elegans, 196 
Trigonomonas, 166 
Triioculina, 184, 185 
Trimastigaceae, 58 
Trimastigida, 58, 165 
Trimastigidae, 58 
Trimastix, 58 
Trinema, 191 
Triplagia, 198 
Triposolenia, 103 
Triposolenia Ambulatrix, 104 
Tripylaria, 199 
Tripylea, l99 
Tripyleen, 199 
Tripylina, 199 
Triticina, 188 
Trochammina, 186 
Trochamminida, 186 
Trochamminidae, 186 
Trochamminina, 186 
Truffles, 135 
Tryblidacea, 134 
Tryblidaceae, 134 
Tryblidieae, l33 
Trypanophidae, 161 
Trypanophis, 161 
Trypanoplasma, 161 
Trypanoplasmida, 159, 161 
Trypanoplasmidae, 161 
Trypanosoma, 162 
Trypanosoma Brucii, 160, 162 
Trypanosoma Cruzi, 162 
Trypanosoma equinum, 162 
Trypanosoma equiperdum, 162 
Trypanosoma Evansi, 162 
Trypanosoma gambiense, 162 
Trypanosoma Lewisi, 160 
Trypanosomata, 158 
Trypanosomatidae, 161 
Trypanosomes, 161, 212 
Trypanosomidae, 161 
Trypanosomidea, 158 
Tuberacea, 135 
Tuberaceae, 134 
Tuberales, 134 
Tuberculariaceae, 141 
Tuberculariea, l4l 
Tubercularieae, 141 
Tubercularini, 141 
Tuberineae, 134 

Tubifer, 175 
Tubiferaceae, 175 
Tubiferida, 174, 175 
Tubiferidae, 175 
Tubinella, 185 
Tubulina, 175 
Tubulinaceae, 175 
Tubulinidae, 175 
Tuburcinia, 149 
Tulasnella, 149, 150 
Tulasnella sphaerospora, 
Tulasnellales, 149 
Tulostoma, 155 
Tulostomataceae, 155 
Tulostomea, 155 
Tulostomei, 155 
Tunicates, 216 
Turillina, 188 
Turkeys, 210 
Turtles, 211 
Tuscarilla, 200 
Tuscarora, 200 
Tuscarorida, 200 

Ulvina aceti, 24 
Umbina aceti, 24 
Uncinula, 132, 133 
Uniflagellatae, 110 
Urceolaria, 235 
Urceolaridae, 235 
Urceolarina, 235 
Urceolus, 109 
Uredinacea, 148 
Uredinaceae, 148 
Uredinales, 145, 147 
Uredineae, 147 
Uredinees, 147 
Uredo, 147 
Uredo linearis, 147 
Urnula, 235 
Urocentridae, 230 
Urocentrina, 230 
Urocentrum, 230 
Uroglena, 59 
Uroglenopsis, 59 
Uroieptus, 233 
Uromyces, 143 
Urophagus, 166 
Urophlyctis, 117 
Urospora, 216 
Urosporida, 216 
Urosporidae, 216 
Urosporidium, 218 
Urostyla, 233 
Urostylida, 233 
Urostylidae, 233 
Ustilaginacea, 149 
Ustilaginaceae, 149 
Ustilaginales, 149 
Ustilaginea, 146, 149 
Ustilagineae, 149 



The Classification of Lower Organisms 

Ustilago, 149 
Ustilago Heufleri, 145 
Ustilago Hordei, 145 
Uterini, 134, 137 
Uvella, 59 
Uvellina, 188 
Uvigerina, 188 
Uvigerinlda, 188 
Uvigerinidae, 188 

Vacuolaria, 65, 109 

Vacuolaria viridis, 108 

Vacuolariaceae, 109 

Vaginicola, 233 

Vaginifera, 233 

Vaginulina, 187 

Vahlkampfia, 202, 203 

Valsa, 139 

Valvulina, 186 

Valvulinidae, 186 

Vampyrella, 118, 189, 191, 192 

Vampyrellacea, 191 

Vampyrellaceae, 191 

Vampyrelleae, 191 

Vampyrellidae, 191 

Vampyrellidea, 190 

Vaucheria, 67, 76 

Vaucheria Gardner!, 64 

Vaucheria sessilis, 64 

Vaucheriacea, 57, 63, 64 

Vaucheriaceae, 63, 67 

Vaucheriales, 63 

Vaucherioideae, 55 

Venturia, 139 

Venturia inaequalis, 139 

Verbeekina, 188 

Vermes, 9 

Verneulina, 186 

Verneulinidae, 186 

Veronica, 69 

Verrucariacea, 139 

Vertebralina, 184, 185 

Vertebrates, 161, 165, 166, 167, 210, 211 

Vibrio, 23 

Vibrio Protheus, 201 

Virgulina, 188 

Volvox Chaos, 201 

Vorticclla, 223, 226, 233, 235 

Vorticellidae, 233 

Vorticellina, 233 

Vorticialcs, 179 

Vorticialis, 186, 187 

Vulvulina, 186 

Wagnerella, 193, 194 
Wardia, 221 

Water molds, 77 

Whales, 71 
Wheat, 148 
Wood roach, 166, 169 
Worms, 215, 217, 220 
Worms, annelid, 216, 219, 221 
Worms, oligochaet, 222 
Worms, polychaet, 211 
Worms, siphunculid, 210 
Woronina, 179 
Woroninaceae, 179 
Woroninidae, 179 
Wrangelieae, 47 

Xanthomonadina, 63 
Xanthomonas, 23 
Xenococcus, 36 
Xiphacantha, 197 
Xylaria, 139 

Zanardinia, 88 

Zea Mays, 6 

Zelleriella, 229 

Zonaria, 87 

Zooflagellata, 157 

Zoomastigina, 157 

Zoomastigoda, 157, 178 

Zoomastigophorea, 157 

Zoopagacea, 123, 124 

Zoopagaceae, 124 

Zoopagales, 121 

Zoopage, 124 

Zoophagus, 81 

Zoosporidae, 191 

Zoosporidea, 191 

Zoosporidia, 190 

Zoothamnium, 233 

Zooxanthellae, 194 

Zostera, 203 

Zschokkella, 221 

Zygochytrium, 118 

Zygocystis, 216 

Zygocystida, 216 

Zygocystidae, 216 

Zygomyceteae, 121 

Zygomyceten, 121 

Zygomycetes, 76, 1 18, 120, 121, 122, 127, 

Zygophyceac, 53 
Zygophyta, 53 
Zygorhynchus, 124 
Zygostephanus, 198 
Zythiacea, 141 
Zythiaceac, 141